Usefulness of the Umbrella Species Concept as a Conservation Tool

Article (PDF Available)inConservation Biology 18(1):76-85 · February 2004with 2,996 Reads 
How we measure 'reads'
A 'read' is counted each time someone views a publication summary (such as the title, abstract, and list of authors), clicks on a figure, or views or downloads the full-text. Learn more
DOI: 10.1111/j.1523-1739.2004.00450.x
Cite this publication
FULL-TEXT OF THE WHOLE PAPER AVAILABLE AT: ABSTRACT: In the face of limited funding, knowledge, and time for action, conservation efforts often rely on shortcuts for the maintenance of biodiversity. The umbrella species concept - proposed as a way to use species requirements as a basis for conservation planning-has recently received growing attention. We reviewed the literature to evaluate the concept's general usefulness. An umbrella species is defined as a species whose conservation is expected to confer protection to a large number of naturally co-occurring species. This concept has been proposed as a tool for determining the minimum size for conservation areas, selecting sites to be included in reserve networks, and setting minimum standards for the composition, structure, and processes of ecosystems. Among the species suggested as potential umbrellas, most are large mammals and birds, but invertebrates are increasingly being considered. Eighteen research papers, most of which were based on hypothetical reserves or conservation networks, have provided evaluations of umbrella species schemes. These show that single-species umbrellas cannot ensure the conservation of all co-occurring species because some species are inevitably limited by ecological factors that are not relevant to the umbrella species. Moreover, they provide evidence that umbrella species from a given higher taxon may not necessarily confer protection to assemblages from other taxa. On the other hand, multi-species strategies based on systematic selection procedures (e. g., the focal species approach) offer more compelling evidence of the usefulness of the concept. Evaluations of umbrella species schemes could be improved by including measures of population viability and data from many years, as well as by comparing the efficiency of the proposed scheme with alternative management strategies.
Usefulness of the Umbrella Species Concept
as a Conservation Tool
Grims¨o Wildlife Research Station, Department of Conservation Biology, Forest Faculty, Swedish University
of Agricultural Sciences (SLU), SE-730 91 Riddarhyttan, Sweden
†Department of Natural Sciences, Centre for Landscape Ecology, ¨
Orebro University, SE-701 82 ¨
Orebro, Sweden
Abstract: In the face of limited funding, knowledge, and time for action, conservation efforts often rely on
shortcuts for the maintenance of biodiversity. The umbrella species concept—proposed as a way to use species
requirements as a basis for conservation planning—has recently received growing attention. We reviewed the
literature to evaluate the concept’s general usefulness. An umbrella species is defined as a species whose con-
servation is expected to confer protection to a large number of naturally co-occurring species. This concept
has been proposed as a tool for determining the minimum size for conservation areas, selecting sites to be
included in reserve networks, and setting minimum standards for the composition, structure, and processes
of ecosystems. Among the species suggested as potential umbrellas, most are large mammals and birds, but
invertebrates are increasingly being considered. Eighteen research papers, most of which were based on hypo-
thetical reserves or conservation networks, have provided evaluations of umbrella species schemes. These show
that single-species umbrellas cannot ensure the conservation of all co-occurring species because some species
are inevitably limited by ecological factors that are not relevant to the umbrella species. Moreover, they provide
evidence that umbrella species from a given higher taxon may not necessarily confer protection to assemblages
from other taxa. On the other hand, multi-species strategies based on systematic selection procedures (e.g., the
focal species approach) offer more compelling evidence of the usefulness of the concept. Evaluations of um-
brella species schemes could be improved by including measures of population viability and data from many
years, as well as by comparing the efficiency of the proposed scheme with alternative management strategies.
Utilidad del Concepto de Especie Paraguas como Herramienta de Conservaci´on
Resumen: Ante la escasez de financiamiento, de conocimiento y tiempo para actuar, los esfuerzos de con-
on a menudo conf´
ıan en m´
etodos r´
apidos para el mantenimiento de la biodiversidad. El concepto
de especie paraguas - propuesto como una manera de utilizar los requerimientos de las especies como base
para la planeaci´
on de conservaci´
on – ha recibido mayor atenci´
on recientemente. Revisamos la literatura para
evaluar la utilidad general del concepto. Una especie paraguas se define como una especie cuya conservaci´
ıa protecci´
umero de especies que coexisten naturalmente. Este concepto se ha propuesto
como una herramienta para determinar el tama˜
no m´
ınimo de ´
areas de conservaci´
on, seleccionar sitios a
incluir en redes de reservas y para fijar las normas m´
ınimas para la composici´
on, estructura y procesos de los
ecosistemas. La mayor´
ıa de las especies que se sugieren como paraguas potenciales son mam´
ıferos y aves may-
ores, pero se est´
an considerando a los invertebrados cada vez m´
as. Dieciocho art´
ıculos cient´
ıficos, la mayor´
basados en reservas o redes de conservaci´
on hipot´
eticas, han proporcionado evaluaciones de proyectos con
especies paraguas. Estos muestran que una sola especie paraguas no puede asegurar la conservaci´
on de todas
las especies coexistentes porque algunas especies inevitablemente est´
an limitadas por factores ecol´
ogicos que
no son relevantes para la especie paraguas. M´
as aun, proporcionan pruebas de que especies paraguas de un
on alto determinado necesariamente no confiere protecci´
on a los conjuntos de otros taxones. Por otro lado,
estrategias multiespec´
ıficas basadas en procedimientos de selecci´
on sistem´
aticos (por ejemplo, el m´
etodo de la
especie focal) ofrecen evidencia m´
as convincente de la utilidad del concepto. Se debe mejorar la evaluaci´
Paper submitted October 23, 2002; revised manuscript accepted April 21, 2003.
Conservation Biology, Pages 76–85
Volume 18, No. 1, February 2004
Roberge & Angelstam Umbrella Species and Conservation 77
de los proyectos de especies paraguas incluyendo medidas de viabilidad poblacional, datos de muchos a˜
ıcomo comparando la eficiencia del proyecto propuesto con estrategias de manejo alternativas.
Following the emergence of the concept of biological
diversity, conservation biologists showed growing inter-
est in developing shortcuts to the conservation of whole
biota. The umbrella species concept has been proposed
as a way to use species requirements to guide ecosystem
management. Its main premise is that the requirements
of demanding species encapsulate those of many co-
occurring, less demanding species (Lambeck 1997). By
directing management efforts toward the requirements
of the most exigent species, one is likely to address the
requirements of many cohabitants that use the same habi-
tat. The umbrella species concept is intuitively appealing
and offers a simple, ecologically based shortcut for the
management of communities. Yet the assumption that
providing for the requirements of a few species results
in the protection of most species present in the same
area rarely has been addressed rigorously. Indeed, many
authors have recently made a plea for the empirical valida-
tion of this concept (Caro & ODoherty 1999; Simberloff
1999; Angelstam et al. 2001; Fleishman et al. 2001).
Although the umbrella species concept has been dis-
cussed in the biological conservation literature, no com-
prehensive review has been made that allows assessment
of its general utility as a conservation tool. Some authors
have discussed this concept more thoroughly than others
(e.g., Simberloff 1998; Caro & ODoherty 1999; Zacharias
& Roff 2001; Caro 2003), but their accountsalthough
rich in insightdo not cover all its variants or extensively
examine the empirical evidence for its utility. We sought
to (1) review the history of the umbrella species concept
and distinguish its different uses, (2) evaluate the extent
to which the concept has been validated empirically, and
(3) suggest directions for using umbrella species in con-
servation planning.
History and Uses of the Umbrella Species Concept
The exact origin of the umbrella species concept is neb-
ulous. To our knowledge, Frankel and Soul´e (1981) were
the first to use the term umbrella to suggest that con-
servation measures directed at the largest species could
confer protection to what they called denser species.A
few years later Wilcox (1984) proposed that management
focus on those species whose habitat requirements are
at least as comprehensive as that of the rest of the com-
munity,thus providing a protective umbrellafor other
species. Other authors had put forward the same basic
idea before that, although without using the term um-
brella (Eisenberg 1980; East 1981; Mealy & Horn 1981).
Therefore, this concept was probably used implicitly be-
fore it was defined (Caro 2003). The biological conserva-
tion literature from the 1980s includes a number of ref-
erences to the umbrella species concept (e.g., Ehrlich &
Murphy 1987; Murphy 1988; Peterson 1988), but most of
these papers only state the theoretical appeal of that con-
cept without evaluating its validity. Since the beginning
of the 1990s, attention to the umbrella species approach
has grown. Instead of merely discussing the appeal of
this idea, many researchers have critically examined the
potential usefulness of proposed umbrella taxa for man-
agement (e.g., Ryti 1992; Beier 1993; Launer & Murphy
1994; Berger 1997; New 1997; Fleury et al. 1998), and
others have suggested ecologically based methods or cri-
teria for the selection of umbrella species (e.g., Lambeck
1997, 1999; Caro & ODoherty 1999; Fleishman et al.
2000, 2001).
A large number of definitions have been proposed for
umbrella species that emphasize different uses and prop-
erties (Zacharias & Roff 2001). This has resulted in some
confusion about the meaning of the term. Fleishman et al.
(2000) recently suggested a broadly applicable definition
for an umbrella species: a species whose conservation
confers a protective umbrella to numerous co-occurring
species.This definition is somewhat tautological, how-
ever, because it includes the term umbrella. A potentially
consensual alternative could be a species whose conser-
vation confers protection to a large number of naturally
co-occurring species. We use this general definition as a
basis throughout this review and define narrower vari-
ants of the concept in context-specific applications. We
use the phrase umbrella scheme to describe any pro-
posed conservation plan based on one or several umbrella
species. The co-occurring species that are likely to be pro-
tected by conservation activities directed at the umbrella
species are hereafter referred to as beneficiary species.
Several more or less overlapping subconcepts of umbrella
species can be distinguished, all of which conform to our
general definition.
Umbrella Species with Large Area Requirements
In its classic form and at the local scale, the umbrella
species concept refers to the minimum area requirements
of a population of a wide-ranging species (Wilcox 1984).
Conservation Biology
Volume 18, No. 1, February 2004
78 Umbrella Species and Conservation Roberge & Angelstam
It hinges on the assumption that providing enough space
for species with large area requirements will also shelter
a whole suite of species with more modest spatial needs.
Because organisms with a large body size also tend to
have large home ranges (McNab 1963), maintaining vi-
able populations of those species requires the conserva-
tion of large tracts of habitat. For that reason, large-bodied
organisms have been favored as prospective umbrellas.
Species in this category are usually vertebrates, typically
large mammalian carnivores (Eisenberg 1980; East 1981;
Peterson 1988; Shafer 1990; Noss et al. 1996; Carroll et al.
2001), herbivores (Mealy & Horn 1981; Wallis de Vries
1995; Berger 1997), or birds (Martikainen et al. 1998;
Suter et al. 2002).
Umbrella Species Used to Select Sites
In addition to its use in determining minimum area for
protection, the umbrella species concept has been pro-
posed as a tool for defining reserve networks at larger
geographic scales. In the site-selection umbrella concept,
the occurrence of single- or multi-species umbrellas, or,
alternatively, the species richness of an umbrella taxon, is
used as a basis for the selection of siteshabitat patches,
ecosystems, planning units, or even large grid squares
to be preserved in a conservation network. The general
methodology consists of setting aside for protection the
totality or a given proportion of the sites where the um-
brella(s) occur(s) or where species diversity for the um-
brella taxon is high (Kerr 1997). It is presumed that pop-
ulations of a number of beneficiary species should be
protected in the network (Ryti 1992; Launer & Murphy
1994; Kerr 1997; Fleishman et al. 2000, 2001).
The Extended Umbrella Concept
The umbrella species concept builds on the assumption
that the requirements of demanding species should en-
compass those of many other species (Lambeck 1997).
This assumption is stated explicitly for area needs in the
subconcept of area-demanding umbrella species. But why
should this tool be restricted to area requirements? In a
variant that we would like to call the extended-umbrella
species concept, the basic idea is broadened to include
other attributes of landscapes such as habitat connectiv-
ity, the occurrence of various ecosystem processes, or the
distribution of scarce resources. For example, Fleury et al.
(1998) and van Langevelde et al. (2000) used, respectively,
the California Gnatcatcher (Polioptila californica) and
the European Nuthatch (Sitta europaea) as umbrella bird
species for reserve network planning. Instead of using
only the occurrences or area requirements of these birds
for reserve site selection, they also considered their need
for connectivity. Fleury et al. (1998) did so by simulating
movement corridors to link subpopulations, whereas van
Langevelde et al. (2000) used threshold between-patch
distances for allowing dispersal in the umbrella species.
The underlying assumption here is that landscapes that
are sufficiently connected for the umbrella species should
also be functional for many other species.
Lambeck (1997) gave new life to the umbrella species
idea by extending it to a large range of factors that threaten
species persistence in managed landscapes. He proposed
to identify a suite of focal speciesthat would be used
to define the spatial, compositional, and functional at-
tributes that must be present in a landscape. Lambecks
process involves identifying the main threats and select-
ing focal species among the species that are most sensi-
tive to each threat. The requirements of those species
would then be used to guide conservation or restora-
tion. Thus, the focal species approach is consistent with
the general definition of umbrella species. It simply goes
beyond earlier approaches by addressing habitat quality
and by proposing systematic criteria for the selection of
a suite of umbrella (focal) species. Lambeck (1997)
suggests four threat categories for focal species: area-
limited, resource-limited, dispersal-limited, or process-
limited. Each of these categories should be represented
by one or a few focal species.
By reviewing the conservation biological literature on
the umbrella species concept, we assessed the relative
prevalence of (1) the main variants of the concept and
(2) the different taxa proposed as umbrellas. We exam-
ined biological database information and reference lists
from relevant papers. We only considered papers pub-
lished in English. In this review we included 110 peer-
reviewed articles, book chapters, and papers from con-
ference proceedings that discussed the umbrella species
concept. Papers dealing with this concept without using
the term umbrella were included if they complied with
one of the three variants of the concept described above.
Because we addressed an applied science issue, we also
considered published governmental research papers and
technical reports. We evaluated papers that dealt with
one or more of the three main umbrella categories (Ta-
ble 1). Not surprisingly, mammals, birds, and to a lesser
extentinsects dominate as suggested or evaluated um-
brella species. Although mammals have often been pro-
posed as area-demanding umbrellas, birds and insects
seem to have been preferred for the site-selection and
extended-umbrella concepts.
Evaluating the Umbrella Species Concept
If the umbrella species concept is to be promoted from an
interesting management hypothesis or social hook (Lin-
denmayer & Fischer 2003) to an actual conservation tool,
Conservation Biology
Volume 18, No. 1, February 2004
Roberge & Angelstam Umbrella Species and Conservation 79
Table 1. The main variants of the umbrella species concept in the biological conservation literature.
Number of papers suggesting or evaluating different taxa
Total no. of papers
Umbrella species category Field of application discussing concept mammals birds herptiles fish invertebrates plants no suggestion or evaluation
Area-demanding umbrella setting the minimum size for
a nature reserve
56 23 6 1 0 0 1 29
Site-selection umbrella prioritizing of patches to be
included in a reserve
15 4 7 1 1 6 3 2
Extended umbrella concept,
including the focal-species
setting minimum values for
landscape composition,
configuration, resources, or
48 8 16 3 1 12 1 24
All categories 110 31 26 5 2 18 4 51
Based on 110 reviewed papers. Rows and columns are not additive because one paper may discuss several umbrella categories and suggest or evaluate species from several taxa.
it is crucial that the approach be validated empirically. A
common remark is that the umbrella species concept has
rarely if ever been tested (Simberloff 1999). Contrary to
classic scientific hypotheses, the umbrella species con-
cept is not subject to falsification (Lindenmayer et al.
2002). Obviously, lack of support for the validity of the
concept in certain conditions does not constitute a proof
of the general invalidity of the tool. Moreover, the results
of such testsusually cannot be classified simply as suc-
cesses or failures in a binary manner, but must rather be
evaluated according to some quantitative measure.
To be considered for empirical evaluation of the um-
brella species concept (Table 2), a study had to fulfill two
criteria. First, it had to present an evaluation of a hypo-
thetical or actual conservation scheme (e.g., spatially de-
limited reserve or network) based on an umbrella species.
Merely describing general patterns of co-occurrence over
an entire region provides information on the indicator
value of the different species but is not sufficient for con-
stituting an evaluation of the umbrella function. Second,
the study had to provide a quantitative measure of the
level of protection conferred to beneficiary species, such
as the number of species represented in the conservation
network or the probability of beneficiary species main-
taining viable populations. Among the 110 research pa-
pers we included in this review, only 18 evaluated the
performance of an umbrella species scheme according to
these criteria.
Evaluations of Area-Demanding Umbrella Species
Most empirical evaluations of the umbrella species con-
cept based on area requirements suggest that it has a lim-
ited effectiveness (Table 2). Noss et al. (1996) investigated
the umbrella value of proposed grizzly bear (Ursus arc-
tos) recovery zones by looking at how many species of
different taxa would have more than 10% of their state-
wide distribution captured by the conservation plan (i.e.,
a surrogate for population viability). Although birds, mam-
mals, and amphibians would be well covered, other taxa
such as reptiles would be underrepresented. A recent
study by Carroll et al. (2001) based on modeling habitat
suitability for the grizzly bear, fisher (Martes pennanti),
lynx (Lynx canadensis), and wolverine (Gulo gulo) il-
lustrated the need for a multi-species approach if those
carnivores are to be used as umbrella species. Unfortu-
nately, these authors only assessed the overlap in prior-
ity areas among those four carnivores and did not evalu-
ate the umbrella function for other species. In an evalua-
tion of the black rhinoceros (Diceros bicornis)asanum-
brella species, Berger (1997) used a different approach.
He examined the umbrella value of the rhinoceros for six
large herbivore species by calculating the frequency with
which the mean population size for each species would
be maintained at a given level within the area needed by
the rhinoceros population. Annual differences in rainfall
Conservation Biology
Volume 18, No. 1, February 2004
80 Umbrella Species and Conservation Roberge & Angelstam
Table 2. Evaluations of the usefulness of different types of umbrella species for conservation planning.
Umbrella species Investigated umbrella
category and citation Ecosystem type taxon/taxaaTargeted beneficiary taxa Protection conferredb
Area-demanding umbrella
Noss et al. 1996 Rocky Mountains, northern
grizzly bear, Ursus arctos
amphibians, reptiles, birds,
mammals, and plant
Berger 1997 African desert black rhinoceros, Diceros
bicornis (S)
six large herbivore species limited
Martikainen et al.
boreal forest, Fennoscandia White-backed
Dendrocopos leucotos
threatened saproxylic
Andelman & Fagan
diverse regions of the
continental U.S.
large carnivores of concern
species of concernfrom
various taxa
limited to ineffective,
depending on databases
and on surrogate scheme
Andelman & Fagan
diverse regions of the
continental U.S.
most widespread species
of concern, from diverse
taxa (M)
species of concernfrom
various taxa
limited to ineffective,
depending on databases
and on surrogate scheme
Caro 2001 East African deciduous
forests, floodplains, and
large mammals (M) small, nonvolant mammals ineffective
Suter et al. 2002 alpine coniferous forests,
Capercaillie, Tetrao
urogallus (S)
birds effective for red-listed
mountain birds;
ineffective for bird
diversity in general
Caro 2003 East African deciduous
large mammals (M) large and middle-sized
mammals, small
carnivores, small native
rodents and insectivores
effective for large and
middle-sized mammals as
well as small carnivores;
ineffective for rodents
and insectivores
Site-selection umbrella
Murphy & Wilcox
high-elevation boreal
habitats separated by
arid scrub; canyon
riparian habitats,
western North America
birds and mammals (M) butterflies limited to effective,
depending on spatial
Ryti 1992 islands and canyons of
southwestern U.S.
plants and birds (M) plants, birds, mammals,
and reptiles
limited (birds) to effective
Launer & Murphy
serpentine soil-based
grasslands, southwestern
bay checkerspot butterfly,
Euphydryas editha
bayensis (S)
plants limited
Kerr 1997 North America north of
mammalian carnivores (M) invertebrates ineffective
Andelman & Fagan
diverse regions of the
continental U.S.
habitat specialists of
concern, from diverse
taxa (M)
species of concernfrom
various taxa
limited to ineffective,
depending on databases
and on surrogate scheme
Fleishman et al.
mountain canyons in the
Great Basin, western U.S.
butterflies (M) butterflies effective
Fleishman et al.
coastal chaparral
shrubland; eastern
broadleaf forest; xeric
shrub steppe, North
butterflies and birds (M) butterflies and birds effective within taxa;
limited across taxa
Poiani et al. 2001 mixture of grassland and
forest, northern U.S.
Greater Prairie Chicken,
Tympanuchus cupido
rare plants and animals,
natural communities
Rubinoff 2001 coastal sage scrub,
southwestern U.S.
California Gnatcatcher,
Polioptila californica (S)
three specialized insect
species (Lepidoptera)
Bonn et al. 2002 southern Africa threatened and/or endemic
birds (M)
birds limited
Extended umbrella species concept
Fleury et al. 1998 coastal sage scrub,
southwestern U.S.
California Gnatcatcher (S) plant and animal species of
Watson et al. 2001 temperate woodland
remnants, southeastern
Hooded Robin,
Melanodryas cucullata,
and Yellow Robin,
Eopsaltria australis (M)
woodland birds effective with ideal
guidelines; limited with
less stringent guidelines
adapted to the social
aAbbreviations: S, single-species umbrella; M, multispecies umbrella.
bInterpretation (by the authors of this review) of the original authorsconclusions. The protection conferred by the umbrella scheme was
summarized in three classes: effective, limited, and ineffective.
Conservation Biology
Volume 18, No. 1, February 2004
Roberge & Angelstam Umbrella Species and Conservation 81
caused fluctuations in population size and movements
for the six species, but the rhinoceros populations did
not vary accordingly. Berger (1997) concluded that the
population of 28 rhinoceroses was unlikely to assure the
populations of the other species in excess of 250 individ-
In East Africa, Caro (2001, 2003) observed that pop-
ulations of edible ungulates and small carnivores were
lower outside than inside a national park designed for
large mammals. On the other hand, average species diver-
sity and abundance of small mammals were greater out-
side than inside the reserve. Based on these observations
he concluded that delineating a reserve using umbrella
species may benefit populations of some background taxa
but by no means all. As discussed by Caro (2001), in such
studies it is crucial to consider the identity of the potential
beneficiary species in addition to their richness and abun-
dance. Attention should be given to the species of con-
cern that are sensitive to human disturbances (e.g., Fleury
et al. 1998; Lambeck 1999; Andelman & Fagan 2000). In
an assessment of the potential of the Capercaillie (Tetrao
urogallus) as an umbrella species, Suter et al. (2002) ex-
amined separately the umbrella effect for all bird species
and for mountain species of concern included in the Swiss
Red List. The distribution of the capercaillie was related
to high species richness in mountain birds of concern,
but this was not the case when ubiquitous bird species
were included in the analyses.
Suter et al.s (2002) study provided one of the few ex-
amples of substantial support for the concept of area-
demanding umbrella species. This is explained by the
fact that they considered variations in vegetation struc-
ture in their study area and addressed the question of
whether the potential beneficiary species had, as a group,
specialized habitat requirements similar to those of the
umbrella species. Similarly, Martikainen et al. (1998) pro-
vided evidence for the potential umbrella value of the
White-backed Woodpecker (Dendrocopos leucotos)for
threatened beetles dependent on the same resources
that is, decaying deciduous wood within successional
deciduous forest. Contrary to the approaches of Suter
et al. (2002) and Martikainen et al. (1998), however, most
other approaches to the concept of area-demanding um-
brella species have been based coarsely on the area re-
quirements of the purported umbrella species without
attention to the habitat needs of the potential beneficiary
Evaluations of Site-Selection Umbrella Species
The different evaluations of the site-selection umbrella
concept have yielded varied results (Table 2). Murphy
and Wilcox (1986) demonstrated that the validity of
vertebrate-based management for maintaining butterfly
diversity is scale dependent. They concluded that re-
gional management of vertebrates would probably pro-
vide for the requirements of invertebrates, but that inten-
sive fragmentation on a local scale may cause declines in
invertebrates without affecting vertebrates. Kerr (1997),
however, provided evidence that a hypothetical system
of large-scale reserves designed to include all carnivore
species in a minimum of sites would not provide good pro-
tection for invertebrates. For gulf islands and canyons in
California, Ryti (1992) found that a reserve network con-
taining at least one occurrence of all plant species found in
the regional pool covered vertebrates well, whereas such
a network for birds did not provide equivalent protection
for other taxa. However, the plant network required set-
ting aside an area much larger than the area needed for
the bird network. Thus, the relative performance of these
two taxa as umbrellas surely would have been different
had issues of efficiency been considered.
Using occurrence data for species of concern from a
variety of taxa, Andelman and Fagan (2000) evaluated a
variety of site-selection umbrella schemes and compared
their efficiency to that of random sets of species. Only
a few umbrella schemes proved more efficient than ran-
dom schemes made up of similar numbers of species. On
this basis, they concluded that there is little evidence sup-
porting the utility of umbrella species. One limitation of
their study is that they automatically included in a given
surrogate scheme all species in the databases that fulfilled
the selection criteria for that scheme. For example, their
habitat specialistscheme was based on the occurrences
of all habitat specialist species found in the databases, re-
sulting in the need to set aside a very large proportion
of the sites. Many such schemes performed well in terms
of species protection but were excessively costly (and
thereby less efficient) because they required too many
conservation areas. Hence, these authors showed that
multi-species schemes might fail to provide an efficient
management alternative if the suite of umbrella species
were selected without addressing issues of parsimony or
complementarity. In Southern Africa, Bonn et al. (2002)
simulated complementary networks representing threat-
ened and endemic birds with the minimum possible num-
ber of conservation areas and assessed whether many
other bird species would be conserved in these areas.
These networks performed better than random schemes,
although they did not guarantee the representation of
overall species diversity.
Fleishman et al. (2001) evaluated different site-selec-
tion umbrella strategies of butterflies and birds and com-
pared their efficiency to that of random schemes. They
ranked species according to their value as umbrellas based
on three criteria: medium rarity, sensitivity to human dis-
turbance, and percentage of co-occurring species (Fleish-
man et al. 2000). Their multispecies umbrella schemes
proved to be efficient conservation tools that allowed the
area to be set aside for achieving a given level of species
protection to be minimized. Launer and Murphy (1994),
Poiani et al. (2001), and Rubinoff (2001) assessed the
Conservation Biology
Volume 18, No. 1, February 2004
82 Umbrella Species and Conservation Roberge & Angelstam
validity of a butterfly, a tetraonid bird, and a passerine
bird as single-species umbrellas for plants, rare animals
and plants, and invertebrate species, respectively (Table
2). They came to a common conclusion: single species are
unlikely to function as effective site-selection umbrellas
for the protection of other taxonomic groups.
Most of those researchers used the occurrences of the
umbrella species as a basis for reserve site selection and
assumed complete data sets. It is possible that some of the
recorded absences of the umbrella species were due to
imperfections of the inventory technique. In such cases,
the planner would fail to include some of the sites that
should actually be set aside for protection. On the other
hand, incomplete data sets on the occurrences of the po-
tential beneficiary species would result in an underesti-
mation of the efficiency of the umbrella scheme.
Evaluations of the Extended Umbrella Concept
The extended umbrella concept has rarely been eval-
uated (Table 2). Fleury et al. (1998) designed a hypo-
thetical reserve for the California Gnatcatcher, including
corridors to connect subpopulations and buffer zones.
It afforded only limited protection to red-listed animal
and plant species because some species required local-
ized habitat types or had area needs larger than those
of the gnatcatcher. Thus, this species alone cannot pro-
vide the basis from which to derive minimum standards
for landscape attributes in this coastal sage scrub ecosys-
tem (see also Rubinoff 2001). Similarly, proposed re-
serves for the Northern Spotted Owl (Strix occidentalis)
in the Pacific Northwest were insufficient to conserve
Marbled Murrelets (Brachyramphus marmoratus) and
some aquatic ecosystems (summarized in Franklin 1994;
Wilcove 1994), which is not surprising given that the
owl lives on land. Bonn and Schr¨oder (2001) and Ranius
(2002) partially evaluated the potential of carabid beetles
as umbrella species. Although these researchers provided
evidence for the indicator value of their target beetle
species, they did not (contrary to the studies included in
Table 2) evaluate any hypothetical conservation scheme
based on their prospective umbrella species.
The focal species approach has been only partially val-
idated. The focal species selection process has been ap-
plied in a few case studies in Australia (Lambeck 1999;
Brooker 2002), Italy (Bani et al. 2002), and the United
States (Hess & King 2002), but these investigations did
not include evaluation of the umbrella function of the se-
lected species. In eastern Australia, Watson et al. (2001)
proposed concrete guidelines for conservation planning
based on the needs of sensitive birds and modeled the
probability of occurrence of different species in rem-
nants of various size and habitat complexity. They con-
cluded that their revegetation guidelines would provide
habitat for about 95% of the resident woodland birds in
the region.
Evaluations of the subconcept of area-demanding um-
brella species provide little support for its utility for the
conservation of biodiversity as a whole. Conservation
schemes based uniquely on area requirements will in-
evitably be flawed unless they also incorporate the needs
of sensitive species requiring specialized habitat types.
The site-selection umbrella species concept, on the other
hand, appears more useful. The umbrella value of single
species is generally low because some species are lim-
ited by ecological factors that are not relevant to the
umbrella species. However, rigorous assessments using
multiple species and systematic selection criteria show
that umbrella species can be an effective tool for pri-
oritizing remnants to be included in conservation net-
works (e.g., Fleishman et al. 2000, 2001). The extended-
umbrella species concept has rarely been evaluated. Lam-
becks (1997) focal species approach has been assessed
only partially, and it remains to be seen whether land-
scapes designed for the most sensitive suite of species
can retain nearly all co-occurring species.
What would constitute a robust, empirical evaluation
of the umbrella species concept? The examples provided
above stress the need to measure the efficiency of the
umbrella schemethat is, to present the results in the
form of a ratio of the benefits (e.g., number of species or
habitat types protected) to the costs (e.g., total area or
number of sites required). This allows for comparison of
the efficiency of the umbrella scheme with that of alter-
native strategies. A number of researchers have looked
at whether the umbrella schemes performed significantly
better than random schemes or null models (Andelman
& Fagan 2000; Fleishman et al. 2001; Poiani et al. 2001).
Although this makes sense from a statistical point of view,
one may wonder whether more ecologically relevant cri-
teria could be used instead. For instance, comparing the
effectiveness of an umbrella scheme with that of measures
based on general ecological principles would be much
more informative from a management perspective (Poiani
et al. 2001). Examples of such measures could be restora-
tion of a nominated proportion of a landscape to create a
variety of patches of the main native land types, planning
for corridors between isolated patches, and restoration of
watercourses and associated riparian vegetation (Hunter
1999; Lindenmayer et al. 2002).
Another limitation of many evaluations of the umbrella
species concept is that they often rely on presence and
absence snapshotdata without distinguishing between
viable or extinction-prone populations (Ryti 1992; Andel-
man & Fagan 2000; Fleishman et al. 2001; but see Berger
1997; Fleury et al. 1998; Bonn et al. 2002). In landscapes
where fragmentation is a relatively recent phenomenon,
some populations in remnants may be sinks or transients
(Watson et al. 2001). Moreover, metapopulation theory
predicts that a number of suitable patches are unoccupied
Conservation Biology
Volume 18, No. 1, February 2004
Roberge & Angelstam Umbrella Species and Conservation 83
at any point in time (Hanski 1999). Therefore, data cov-
ering many years is preferable for assessing the viability
of the (sub)populations and increasing the likelihood of
including all suitable habitat in the conservation network.
Moreover, because many species vary in their habitat re-
quirements at different times of the year, surveys should
ideally cover more than one season.
Finally and most important, none of the studies listed in
Table 2 (except Caro 2001, 2003) provides a direct evalu-
ation of the basic assumption of the umbrella species con-
cept, to show that the conservation measures directed at
the umbrella species actually protect many other species.
Their conclusions are based on hypothetical reserves or
conservation networks derived from current species dis-
tributions, which in turn are dependent on the current
structures of landscapes. Hence, they do not demonstrate
that the implementation of the conservation schemes in
the real world would give the same results. This remains
the greatest challenge in evaluating the umbrella species
concept. In that respect, investigations such as Caros
(2001, 2003) based on data from existing protected ar-
eas and their surroundings are of great interest.
Directions for Using Umbrella Species
in Conservation Planning
A frequent criticism of the umbrella species concept is
that it is improbable that the requirements of one species
would encapsulate those of all other species (Noss et al.
1997; Basset et al. 2001; Hess & King 2002). Evaluations
of the utility of single-species umbrellas have shown con-
vincingly that they do not offer total coverage for whole
communities (Table 2). Hence, there is a need for multi-
species strategies as a means to broaden the width of
the protective umbrella (e.g., Miller et al. 1998; Fleish-
man et al. 2000, 2001; Carroll et al. 2001). Among the
different multi-species approaches, Lambecks (1997) fo-
cal species approach seems the most promising because
it provides a systematic procedure for selection of um-
brella species (Lambeck 1999; Watson et al. 2001; Bani et
al. 2002; Brooker 2002; Hess & King 2002).
What qualities should a dream teamof focal species
possess to be a dependable tool of biodiversity assess-
ment and conservation planning? First, it should cover
the main ecosystem or landscape types of concern in a re-
gion (Angelstam 1998a, 1998b; Hess & King 2002). Here
it might be argued that simply conserving all landscape
types would constitute a more straightforward alterna-
tive. However, given the need to establish concrete and
quantitative landscape design criteria, it is essential to re-
fer to the requirements of the species to know how much
is enough (Hansen et al. 1993; Lambeck 1997, 1999; An-
gelstam et al. 2003). For each landscape type, the most
sensitive group of species in terms of resources, area
requirements, connectivity, and natural processes (e.g.,
fire and flooding regimes) should be selected (Lambeck
1997). Depending on the context of application, the suite
of focal species may need to represent a range of spatial
scales from bioregions and landscapes (Lambeck 1999) to
localized habitats and microhabitats (M¨uhlenberg et al.
1991). In forested landscapes, for example, birds and
mammals may include species that are among the most
sensitive to threatening factors operating at landscape and
bioregional scales (Lambeck 1999), whereas species of in-
vertebrates and nonvascular plants could be among the
most sensitive to threats operating at finer spatial scales
(Angelstam 1998a; Ranius 2002).
One shortcoming of the focal species approach lies in
the difficulty of identifying the most sensitive species for
each threat, given that many taxa are still poorly known
(e.g., invertebrates) (Lindenmayer et al. 2002; Linden-
mayer & Fischer 2003). Moreover, there is an obvious
lack of data about requirements that are difficult to study
quantitatively, such as dispersal (Koenig et al. 1996). New
knowledge on habitat thresholds for the persistence of
sensitive species (Fahrig 2001) will contribute to filling
those gaps. In the face of incomplete data, one can use
expert judgment obtained through workshops or surveys
(Lambeck 1999; Hess & King 2002).
Lindenmayer et al. (2002) assert that a ridiculously large
number of species may be needed and that this would ren-
der the focal species approach inefficient. Although this
certainly may be true in ecologically complex landscapes
such as those found in Australia (Clark 1990), develop-
ment of umbrella strategies based on a suite of species
may be attempted in simpler systems, such as boreal re-
gions. Yet it remains that multi-species umbrella strategies
require more data than do classic applications of single
area-demanding umbrellas. In situations where time is lim-
ited, a philosophy of adaptive management should be ad-
vocated (Walters 1986) whereby strategies based on such
multi-species umbrellas are implemented in some case
studies and are later improved or rejected as more knowl-
edge becomes available. In any case, such an approach
would represent a significant conservation shortcut com-
pared with species-by-species management or planning
for habitat re-creation after further destruction.
Species are going extinct at unprecedented rates, even
if we ignore the possibility of a considerable extinction
debt (Pimm et al. 1995; Hanski & Ovaskainen 2002). The
urgency of the situation leaves no room for paralysis by
overanalysis(sensu Carroll & Meffe 1994). It is no longer
legitimate to assert that the umbrella species concept has
not been tested and therefore should not be applied.
Although there is little evidence for the usefulness of
Conservation Biology
Volume 18, No. 1, February 2004
84 Umbrella Species and Conservation Roberge & Angelstam
single-species umbrellas selected only on the basis of their
large area requirements, some multi-species schemes con-
sidering the occurrence of a range of habitat types and
landscape attributes offer promising conservation av-
enues. Using the umbrella species concept could there-
fore provide an appealing and sound approach for rapid
The umbrella species concept is not a panacea. Even
the most sophisticated umbrella schemes probably can-
not guarantee the protection of absolutely all species. The
real issue, however, is whether the umbrella species con-
cept constitutes an effective conservation tool compared
with alternative methods. If used alone, general guide-
lines based on the conservation of the main native land
types would be difficult to implement because they do
not provide clear estimates of how much is needed of
different ecological attributes. On the other hand, the ex-
tended umbrella species concept provides explicit and
quantitative guidelines that could be useful for the as-
sessment of status and trends as well as for conserva-
tion planning. In the face of incomplete knowledge about
species requirements and ecosystem processes, the pre-
cautionary principle advises us to employ a combination
of methods. The umbrella species concept deserves con-
sideration as one of these.
A list of references for the literature summarized in Table 1
is available on request from the authors. We are grate-
ful to H. Andr´en, V.-A. Angers, M. B´elisle, T. M. Caro, L.
Gustafsson, L. Imbeau, G. Jansson, E. Main, G. Mikusin-
ski, M.-A. Villard, and two anonymous reviewers for their
comments on the manuscript. Financial support during
work on this review was provided by scholarships from
the Natural Sciences and Engineering Research Council
of Canada (NSERC) and Helge Axson Johnsons Fund to
J.-M.R., as well as grants from the World Wildlife Fund and
Mistra to P.A.
Literature Cited
Andelman, S. J., and W. F. Fagan. 2000. Umbrellas and flagships: effi-
cient conservation surrogates or expensive mistakes? Proceedings
of the National Academy of Sciences of the United States of America
Angelstam, P. 1998a. Towards a logic for assessing biodiversity in boreal
forests. Pages 301313 in P. Bachmann, M. K¨ohl, and R. P¨aivinen,
editors. Proceedings of the conference on the assessment of biodi-
versity for improved forest planning, held in Monte Verit`a, Switzer-
land. European Forest Institute proceedings 18. Kluwer Academic
Publishers, Dordrecht, The Netherlands.
Angelstam, P. 1998b. Maintaining and restoring biodiversity in European
boreal forests by developing natural disturbance regimes. Journal of
Vegetation Science 9:593602.
Angelstam, P., M. Breuss, and G. Mikusinski. 2001. Toward the assess-
ment of forest biodiversity at the scale of forest management units:
a European landscape perspective. Pages 5974 in A. Franc, O.
Laroussinie, and T. Karjalainen, editors. Proceedings of the confer-
ence on criteria and indicators for sustainable forest management
at the forest management unit level. European Forest Institute pro-
ceedings 38. Gummerus Press, Saarij¨arvi, Finland.
Angelstam, P., et al. 2004. Habitat modelling as a tool for landscape-scale
conservation: a review of parameters for focal forestbirds. Ecological
Bulletins 51: in press.
Bani, L., M. Baietto, L. Bottoni, and R. Massa. 2002. The use of focal
species in designing a habitat network for a lowland area of Lom-
bardy, Italy. Conservation Biology 16:826831.
Basset, Y., E. Charles, D. S. Hammond, and V. K. Brown. 2001. Short-
term effects of canopy openness on insect herbivores in rain forest
in Guyana. Journal of Applied Ecology 38:10451058.
Beier, P. 1993. Determining minimum ha bitat areasand ha bitat corridors
for cougars. Conservation Biology 7:94108.
Berger, J. 1997. Population constraints associated with the use of black
rhinos as an umbrella species for desert herbivores. Conservation
Biology 11:6978.
Bonn, A., and B. Schr¨oder. 2001. Habitat models and their transfer for
single and multi species groups: a case study of carabids in an alluvial
forest. Ecography 24:483496.
Bonn, A., A. S. L. Rodrigues, and K. J. Gaston. 2002. Threatened and
endemic species: are they good indicators of patterns of biodiversity
on a national scale? Ecology Letters 5:733741.
Brooker, L. 2002. The application of focal species knowledge to land-
scape design in agricultural lands using the ecological neighbour-
hood as a template. Landscape and Urban Planning 60:185210.
Caro, T. M. 2001. Species richness and abundance of small mammals
inside and outside an African national park. Biological Conservation
Caro, T. M. 2003. Umbrella species: critique and lessons from East Africa.
Animal Conservation 6:171181.
Caro, T. M., and G. ODoherty. 1999. On the use of surrogate species in
conservation biology. Conservation Biology 13:805814.
Carroll, C. R., and G. K. Meffe. 1994. Management to meet conservation
goals: general principles. Pages 307335 in G. K. Meffe and C. R. Car-
roll, editors. Principles of conservation biology. Sinauer Associates,
Sunderland, Massachusetts.
Carroll, C., R. F. Noss, and P. C. Paquet. 2001. Carnivores as focal species
for conservation planning in the Rocky Mountain Region. Ecological
Applications 11:961980.
Clark, R. L. 1990. Ecological history for environmental management.
Proceedings of the Ecological Society of Australia 16:116.
East, R. 1981. Species-area curves and populations of large mammals in
African savanna reserves. Biological Conservation 21:111126.
Ehrlich, P. R., and D. D. Murphy. 1987. Monitoring populations on rem-
nants of native vegetation. Pages 201210 in D. A. Saunders, G. W.
Arnold, A. A. Burbidge, and A. J. M. Hopkins, editors. Nature con-
servation: the role of remnants of native vegetation. Surrey Beatty &
Sons, Sydney.
Eisenberg, J. F. 1980. The density and biomass of tropical mammals.
Pages 3555 in M. E. Soul´e and B. A. Wilcox, editors. Conservation
biology: an evolutionary-ecological perspective. Sinauer Associates,
Sunderland, Massachusetts.
Fahrig, L. 2001. How much habitat is enough? Biological Conservation
Fleishman, E., D. D. Murphy, and P. F. Brussard. 2000. A new method for
selection of umbrella species for conservation planning. Ecological
Applications 10:569579.
Fleishman, E., R. B. Blair, and D. D. Murphy. 2001. Empirical validation
of a method for umbrella species selection. Ecological Applications
Fleury, S. A., P. J. Mock, and J. F. OLeary. 1998. Is the California gnat-
catcher a good umbrella species? Western Birds 29:453467.
Frankel, O. H., and M. E. Soul´e. 1981. Conservation and evolution. Cam-
bridge University Press, Cambridge, United Kingdom.
Conservation Biology
Volume 18, No. 1, February 2004
Roberge & Angelstam Umbrella Species and Conservation 85
Franklin, J. F. 1994. Preserving biodiversity: species in landscapes: re-
sponse to Tracy and Brussard. Ecological Applications 4:208209.
Hansen, A. J., S. L. Garman, B. Marks, and D.L. Urban. 1993. An approach
for managing vertebrate diversity across multiple-use landscapes.
Ecological Applications 3:481496.
Hanski, I. 1999. Metapopulation ecology. Oxford University Press, Ox-
ford, United Kingdom.
Hanski, I., and O. Ovaskainen. 2002. Extinction debt at extinctionthresh-
old. Conservation Biology 16:666673.
Hess, G. R., and T. J. King. 2002. Planning open spaces for wildlife. I.
Selecting focal species using a Delphi survey approach. Landscape
and Urban Planning 58:2540.
Hunter, M. L., editor. 1999. Maintaining biodiversity in forest ecosys-
tems. Cambridge University Press, Cambridge, United Kingdom.
Kerr, J. T. 1997. Species richness, endemism, and the choice of areas for
conservation. Conservation Biology 11:10941100.
Koenig, V. D., D. van Vuren, and P. N. Hooge. 1996. Detectability, philopa-
try, and the distribution of dispersal distances in vertebrates. Trends
in Ecology & Evolution 11:514517.
Lambeck, R. J. 1997. Focal species: a multi-species umbrella for nature
conservation. Conservation Biology 11:849856.
Lambeck, R. J. 1999. Landscape planning for biodiversity conservation
in agricultural regions: a case study from the wheatbelt of West-
ern Australia. Biodiversity technical paper 2. Environment Australia,
Launer, A. E., and D. D. Murphy. 1994. Umbrella species and the con-
servation of habitat fragments: a case of a threatened butterfly and
a vanishing grassland ecosystem. Biological Conservation 69:145
Lindenmayer, D. B., and J. Fischer. 2003. Sound science or social hook:
a response to Brookers application of the focal species approach.
Landscape and Urban Planning 62:149158.
Lindenmayer, D. B., A. D. Manning, P. L. Smith, H. P. Possingham, J.
Fischer, I. Oliver, and M. A. McCarthy. 2002. The focal-species ap-
proach and landscape restoration: a critique. Conservation Biology
Martikainen, P., L. Kaila, and Y. Haila. 1998. Threatened beetles in White-
backed Woodpecker habitats. Conservation Biology 12:293301.
McNab, B. K. 1963. Bioenergetics and the determination of home range
size. The American Naturalist 97:130140.
Mealy, S. P., and J. R. Horn. 1981. Integrating wildlife habitat objectives
into the forest plan. Transactions of the North American Wildlife
Conference 46:488500.
Miller, B., R. Reading, J. Strittholt, C. Carroll, R. Noss, M. Soul´e, O.
S´anchez, J. Terborgh, D. Brightsmith, T. Cheeseman, and D. Foreman.
1998. Using focal species in the design of nature reserve networks.
Wild Earth 1998/99:8192.
M¨uhlenberg, M., T. Hovestadt, and J. R¨oser. 1991. Are there minimal
areas for animal populations? Pages 227264 in A. Seitz and V.
Loeschcke, editors. Species conservation: a population-biological
approach. Birkh¨auser Verlag, Basel, Switzerland.
Murphy, D. D. 1988. Challenges to biological diversity in urban areas.
Pages 7176 in E. O. Wilson and F. M. Peter, editors. Biodiversity.
National Academy Press, Washington, D.C.
Murphy, D. D., and B. A. Wilcox. 1986. Butterfly diversity in natural
habitat fragments: a test of the validity of vertebate-based manage-
ment. Pages 287292 in J. Verner, M. L. Morrison, and C. J. Ralph,
editors. Wildlife 2000: modelling habitat relationships of terrestrial
vertebrates. University of Wisconsin Press, Madison.
New, T. R. 1997. Are Lepidoptera an effective umbrella groupfor
biodiversity conservation? Journal of Insect Conservation 1:512.
Noss, R. F., H. B. Quigley, M. G. Hornocker, T. Merrill, and P. C. Paquet.
1996. Conservation biology and carnivore conservation in the Rocky
Mountains. Conservation Biology 10:949963.
Noss, R. F., M. A. OConnell, and D. D. Murphy. 1997. The science of
conservation planning: habitat conservation under the Endangered
Species Act. Island Press, Washington, D.C.
Peterson, R. O. 1988. The pit or the pendulum: issues in large carnivore
management in natural ecosystems. Pages 105117 in J. K. Agee
and D. R. Johnson, editors. Ecosystem management for parks and
wilderness. University of Washington Press, Seattle.
Pimm, S. L., G. R. Russel, J. L. Gittleman, and T. M. Brooks. 1995. The
future of biodiversity. Science 269:347350.
Poiani, K. A., M. D. Merril, and K. A. Chapman. 2001. Identifying
conservation-priority areas in a fragmented Minnesota landscape
based on the umbrella species concept and selection of large patches
of natural vegetation. Conservation Biology 15:513522.
Ranius, T. 2002. Osmoderma eremita as an indicator of species richness
of beetles in tree hollows. Biodiversity and Conservation 11:931
Rubinoff, D. 2001. Evaluating the California Gnatcatcher as an umbrella
species for conservation of southern California coastal sage scrub.
Conservation Biology 15:13741383.
Ryti, R. T. 1992. Effect of the focal taxon on the selection of nature
reserves. Ecological Applications 2:404410.
Shafer, C. L. 1990. Nature reserves: island theory and conservation prac-
tice. Smithsonian Institution Press, Washington, D.C.
Simberloff, D. 1998. Flagships, umbrellas, and keystones: is single-
species management pass´e in the landscape era? Biological Con-
servation 83:247257.
Simberloff, D. 1999. The role of science in the preservation of forest
biodiversity. Forest Ecology and Management 115:101111.
Suter, W., R. F. Graf, and R. Hess. 2002. Capercaillie (Tetrao urogallus)
and avian biodiversity: testing the umbrella-species concept. Con-
servation Biology 16:778788.
van Langevelde, F., A. Schotman, F. Claassen, and G. Sparenburg. 2000.
Competing land use in the reserve site selection problem. Landscape
Ecology 15:243256.
Wallis de Vries, M. F. 1995. Large herbivores and the design of large-scale
nature reserves in Western Europe. Conservation Biology 9:2533.
Walters, C. 1986. Adaptive management of renewable resources.
MacMillan, New York.
Watson, J., D. Freudenberger, and D. Paull. 2001. An assessment of the
focal-species approach for conserving birds in variegated landscapes
in southeastern Australia. Conservation Biology 15:13641373.
Wilcove, D. S. 1994. Turning conservation goals into tangible results:
the case of the spotted owl and old-growth forests. Pages 313329
in P. J. in Edwards, R. M. May, and N. R. Webb, editors. Large-scale
ecology and conservation biology. Blackwell Scientific Publications,
Wilcox, B. A. 1984. In situ conservation of genetic resources: determi-
nants of minimum area requirements. Pages639647 in J. A. McNeely
and K. R. Miller, editors. National parks, conservation and develop-
ment: the role of protected areas in sustaining society. Smithsonian
Institution Press, Washington, D.C.
Zacharias, M. A., and J. C. Roff. 2001. Use of focal species in marine
conservation and management: a review and critique. Aquatic Con-
servation: Marine and Freshwater Ecosystems 11:5976.
Conservation Biology
Volume 18, No. 1, February 2004

Supplementary resource

  • ... This suggests that eels have the potential to be an indicator of river-ocean connectivity. This wide distribution of eels is consistent with the umbrella species concept that the presence of species with large area requirements will also encompass a whole suite of species with more limited spatial needs 13,18 . Additionally, the stable isotopic analyses supported their potential as an umbrella species; the TL of eels was greater than three -these values are normally associated with secondary consumers. ...
    Full-text available
    To monitor and manage biodiversity, surrogate species (i.e., indicator, umbrella and flagship species) have been proposed where conservation resources are focused on a limited number of focal organisms. Using data obtained from 78 sites across six rivers in the mainland Japan and the Amami-Oshima Island, we demonstrate that two anguillids – the Japanese eel (Anguilla japonica) and the giant mottled eel (A. marmorata) – can act as surrogate species for conservation of freshwater biodiversity. Anguillid eels were the widest topographically-distributed species ranging from near the mouth to the upper reaches of rivers. Moreover, stable isotopic analyses indicated that eels are likely one of the highest-order predators in freshwater ecosystems. A significant positive relationship was found between the density of eels and the number of other diadromous species collected. However, the optimal models revealed that both the density of eels and the number of other diadromous species were significantly negatively correlated with distance from the river mouth and cumulative height of trans-river structures from the river mouth to each site. This suggests the positive relationship between eel density and number of other diadromous species was indirect and related to river-ocean connectivity. Given their catadromous life-cycle, and global commercial and cultural importance, as a taxa, anguillid eels can act as indicator, umbrella and flagship species, and a comprehensive surrogate for conservation of freshwater biodiversity.
  • ... We have shown that artificial habitats are important for our target species, albeit differences in breeding site selection between A. obstetricans and P. punctatus (both species coexisted in only five sites) imply that management actions need to target each species separately. An important consideration is to what extent A. obstetricans and P. punctatus can act as umbrella species to help protect populations of other taxa (Roberge and Angelstam 2004). Our analysis of cooccurrence patterns shows that other amphibian species in the study area could benefit from actions improving breeding habitat for A. obstetricans and P. punctatus. ...
    Full-text available
    Amphibians in rural landscapes often utilize various types of artificial constructions originally designed for irrigation, livestock supply, or other purposes (e.g., water tanks or cattle troughs) as breeding sites. These sites potentially function as local refugia; however, their importance for amphibian communities has yet to be widely assessed. Here we evaluate the role of artificial constructions in the persistence of amphibian populations in rural areas of central Spain, focusing on two species of conservation concern: the Common Midwife Toad, Alytes obstetricans (Laurenti 1768), and the Parsley Frog, Pelodytes punctatus (Daudin 1802). We surveyed 130 water bodies at 113 localities in an area of 1,450 km 2 during the breeding season of 2018 and documented the type of breeding site, species abundance, amphibian community structure, and any detectable threats. We found non-random patterns of breeding site selection and amphibian species co-occurrence in which A. obstetricans tended to inhabit artificial water tanks with simpler amphibian communities, whereas P. punctatus tended to co-occur in naturalized ponds in abandoned quarries with complex amphibian community structures. We discuss the relevance of artificial breeding sites for the resilience of amphibian populations and propose conservation measures to improve their efficiency in the face of detected threats, including trap effects, alien species, and chytridiomycosis.
  • ... The effectiveness of conservation surrogates is, however, equivocal and context-dependent, with recognition of many putative umbrella and flagship species failing to optimally protect diversity at local scales, by focusing resources on too few species or failing to account for the ecological needs of other threatened species (Roberge and Angelstam 2004;Caro 2010Caro , 2015Joseph et al. 2011). Traditionally, umbrella species have been chosen based on species-specific traits, such as body size, home-range size, or trophic level and diet, without directly verifying ecological suitability or relationships to diversity and abundance of other species, resulting in reduced protection for other areas and species of high conservation priority (Branton and Richardson 2011;Stuber and Fontaine 2018). ...
    Full-text available
    Conservation surrogates, such as umbrella and flagship species, could help focus South Africa’s limited resources for research and management and enhance the conservation gains from marine protected areas (MPAs). Sharks, rays and chimaeras (Chondrichthyes), which are charismatic and ecologically diverse, are potential umbrella candidates, but tests of the ecological suitability of putative marine umbrella species are lacking. Using baited remote underwater video in and around two MPAs in the Western Cape Province, we assessed the potential of chondrichthyans as an umbrella species-complex by quantifying the relationships and co-occurrence patterns between chondrichthyan abundance and diversity and those of other taxa (primarily teleosts and crustaceans). Sites with abundant chondrichthyans, with catsharks or large sharks (>1 m total length), all had significantly greater abundance and diversity of these other taxa, and associations with species of commercial and conservation interest (e.g. roman Chrysoblephus laticeps). Endemic scyliorhinids (notably dark catshark Haploblepharus pictus) and the broadnose sevengill shark Notorynchus cepedianus also had many strong positive co-occurrences (28% and 21% of interactions, respectively). The puffadder catshark H. edwardsii had the highest centrality of any species, denoting its high connectedness to other taxa. Overall, chondrichthyans, especially the dark and puffadder catsharks and the broadnose sevengill shark, show strong potential as an umbrella species-complex in South Africa.
  • ... Saarman et al. (2018) also predicted individual species and fish assemblages to have vastly different responses to human activity under four hypothetical projects within the marine protected areas of California. Inconsistencies in the conservation targets may be explained by a lack of shared ecological traits among species (Branton & Richardson, 2011;Roberge & Angelstam, 2004), or by the fact that combining multiple species with various habitat needs into one fish guild may minimize the perceived consequence of losing specific habitat types. The results of this study emphasize that impact assessments on single-and multi-species conservation targets should not be used interchangeably to inform permitting decisions or to guide the management of proposed habitat alterations. ...
    • To assess the impacts of human activity on fishes and fish habitat, impact assessment tools use single‐ and multi‐species approaches depending on the ecological and socio‐economic objectives. In Canadian aquatic ecosystems, single‐ and multi‐species impact assessments are guided by the Species at Risk Act and Fisheries Act, respectively. Yet, for species protected under the Species at Risk Act, the sparse data often require alternative approaches to risk assessment. • The goal of this study was to evaluate whether a database‐derived multi‐species tool – the Habitat Ecosystem Assessment Tool (HEAT) – can be used for single‐species impact assessments. Using an empirical example of proposed drain maintenance in a tributary of Lake St. Clair, the net loss of suitable habitat was evaluated across six conservation targets, ranging from single species, such as the pugnose shiner (Notropis anogenus) and the yellow perch (Perca flavescens), to the entire fish assemblage. Model outcomes were compared across various habitat suitability indices, spatial resolutions, and environmental habitat layers. • The net loss of suitable habitat varied widely across conservation targets and was greatest for the rare specialist species (pugnose shiner). Single‐species conservation targets were more sensitive to variation in spatial resolution and uncertainty in model input parameters. The results of this study emphasize that single‐ and multi‐species conservation targets should not be considered equal, especially when species differ in abundance and niche breadth. • This study demonstrates the flexibility of HEAT for evaluating potential impacts of human disturbance on fishes and their habitat. Future development of this tool should expand beyond physical habitat, to include other factors relevant to species distribution and survival (e.g. biotic interactions).
  • Article
    L’île de Porquerolles abrite une faune et une flore originales et diversifiées. Alors que certains groupes ont été particulièrement bien étudiés, aucun recensement des Apiformes n’a été réalisé. Les abeilles sont très diversifiées, l’Abeille domestique n’est qu’une espèce parmi les 977 espèces estimées présentes en France métropolitaine. De plus, ce sont des insectes pollinisateurs essentiels tant pour la flore sauvage que cultivée. Afin de pallier le manque de données sur ce groupe, un inventaire des espèces d’abeilles a été réalisé dans les principaux habitats grâce à une campagne de captures mensuelles au filet sur les fleurs et avec des coupelles colorées de février 2007 à janvier 2008. L’île compte au moins 114 espèces d’abeilles sur un territoire d’une dizaine de kilomètres carrés, soit plus de 12 % du nombre d’espèces d’abeilles présentes en France. Parmi ces espèces, 32 sont peu communes et 15 sont rares en France, comme Ammobates punctatus, Colletes eous, Halictus gemmeus, Hoplitis praestans et Colletes collaris, cette dernière figurant au titre d’espèce en danger sur la liste rouge des abeilles d’Europe de l’UICN. Le suivi de ces populations représente donc un enjeu de conservation pour ces espèces à l’échelle nationale, et même mondiale. Les sites anthropisés se sont avérés être les plus riches en abondance et en espèces d’abeilles. Le réseau d’interactions établi dans notre étude montre que la fragilité de certaines populations végétales peut être liée à celle de leurs insectes pollinisateurs qui sont eux-mêmes rares. Or les abeilles sont plus rarement au coeur de mesures de protection que les végétaux. Ainsi, la protection d’espèces végétales peut aider à la préservation des abeilles qui les butinent et qui peuvent contribuer à leur reproduction. La prise en compte des populations d’insectes pollinisateurs devrait aussi être une action en soi pour aider à la conservation des espèces végétales entomophiles rares.
  • Article
    There are many designations people give to species in the context of conservation, some ecological (e.g., umbrella, keystone) and some cultural (e.g., flagship). However, “iconic” is a term with uses in conservation discourse scholars have yet to examine. Environmental nongovernmental organizations (ENGOs) can shape public opinion on conservation issues; hence, understanding their use of the term “iconic species” could help conservation managers assess and respond to stakeholder’s attitudes regarding proposed conservation policies or efforts. This content analysis covered four years’ worth of press releases (n = 203) containing “iconic species” (or an equivalent phrase; e.g., “iconic wildlife”) from six large ENGOs. We examined the species referenced, affiliated terms (including referenced places), press release objectives, and conservation topics. Results indicate that within ENGO communications, the term iconic species is rarely explicitly defined but is generally used to describe species that share taxonomic similarities with the charismatic megafauna of flagship species. However, iconic species appear to represent specific geographic places and are referenced in relation to diverse conservation topics. Thus, iconic species may serve as conduits for communicating regional issues, such as development and species management policies.
  • Thesis
    Dans les Alpes du Nord françaises, les populations de tétras-lyre sont menacées par la fermeture du paysage induite par la colonisation d'espèces ligneuses sur les pâturages abandonnés. Parmi ces espèces, l'aulne vert est considérée comme une menace majeure pour la conservation de l'habitat de reproduction du galliforme. Dans ce contexte, il est nécessaire de mieux connaître les effets de l'organisation spatiale de l'aulne vert sur l'occurrence du tétras-lyre afin de mieux maîtriser cette dynamique d'embuissonnement à l'échelle d'une gestion locale de cet habitat de reproduction. Ainsi, cette thèse s'est attachée dans un premier temps à mettre en évidence que l'occurrence du galliforme peut être prédite à l'aide d'indicateurs spatialisés relatifs à la structure du paysage induite par la distribution de l'aulne vert sur une étendue occupée au sein de son habitat de reproduction. En effet, des données de présence-absence du galliforme issues d'un échantillonnage systématique ont été combinées à une analyse de l'organisation spatiale de l'aulne vert. Ceci a permis d'évaluer l'effet de certains indices paysagers sur la probabilité de présence du galliforme à l'aide de modèles de régression logistique. Nos résultats ont principalement montré que les poules de tétras-lyre sont sensibles à l'organisation spatiale de l'aulne vert alors que ce n'est pas le cas pour les coqs. Plus précisément, le taux de recouvrement de l'aulne vert, le nombre de patchs d'aulne vert et la somme des longueurs de lisières d'aulne vert sont apparus comme étant des prédicteurs significatifs de l'occurrence des poules.Ces trois prédicteurs ont ensuite été utilisés pour construire des modèles spatialement explicites décrivant l'occupation de l'espace par les poules en fonction d'une carte de distribution de l'aulne vert. Après validation des prédictions des modèles, le modèle le plus pertinent et le plus valide pour prédire l'occurrence des poules est celui utilisant la somme des longueurs de lisières associée au taux de recouvrement de l'aulne vert. D'autre part, ce modèle a été combiné à un modèle de simulation de l'expansion de l'aulne vert dans le temps et l'espace à l'aide d'un modèle à automate cellulaire stochastique. Ainsi, nous avons pu simuler l'évolution de la distribution de l'occurrence des poules en fonction de l'évolution des patrons d'organisation du paysage induite par l'expansion de l'aulne vert.La mise en évidence d'un lien entre des patrons paysagers et l'occupation de l'espace par les poules à une échelle fine suggère qu'intégrer des indices paysagers spatialisés dans un modèle spatial et temporel devrait permettre de développer un outil d'aide à la gestion locale de l'habitat du galliforme reposant sur une approche d'écologie du paysage.
  • Thesis
    Dans le contexte du changement global et de la perte de la biodiversité, la prise en compte de cette dernière dans la gestion forestière reste encore très générale (embryonnaire). Les démarches ne sont pas encore opérationnelles alors que les gestionnaires ont besoin d'outils destinés directement à la planification. Ainsi, il est devenu urgent d'améliorer les approches d'évaluation et de suivi de la biodiversité et de développer un cadre conceptuel à partir des indicateurs biologiques qui puissent servir dans la prise de décisions. Ce travail repose sur une approche associant l'enjeu de conservation d'une espèce sauvage patrimoniale dans un paysage de montagne à la maîtrise d'une gestion forestière qui détermine la qualité de son habitat. Dans le cadre concret de la gestion forestière d'un Parc Naturel Régional où les enjeux sont multiples (écologiques, économiques, touristiques), les acteurs concernés s'interrogent moins sur le "comment gérer" que sur le "où gérer" ; en d'autres termes, définir les périmètres de bonne qualité d'habitat est nécessaire afin d'optimiser la gestion à l'échelle optimale. Notre travail est de procurer des outils aux gestionnaires et de poser les jalons scientifiques d'une approche holistique inspirée de l'écologie du paysage. Nous partons ainsi de l'hypothèse selon laquelle les variables environnementales ainsi que la structure du paysage influencent la présence d'une espèce indicatrice de la diversité biologique d'un milieu. Dans ce travail nous nous appuyons sur cette hypothèse pour développer une démarche spatialement explicite qui permettrait d'atteindre l'objectif de conservation dans le cadre d'un processus de gestion forestière adaptative, en étudiant le cas de la gélinotte des bois (Tetrastes bonasia L. 1758) dans le Parc Naturel Régional de Grande Chartreuse (Alpes françaises). Ce modèle prédictif de l'occurrence de la Gélinotte des bois en Chartreuse s'appuie sur des indicateurs environnementaux et paysagers aisément mesurables et spatialisables. Ceci permettra d'établir un diagnostic de la qualité de l'habitat, d'approfondir les connaissances sur les relations entre la structure du paysage et l'espèce à différentes échelles et de déterminer les secteurs d'habitat qui sont les plus appropriés à la Gélinotte des bois en zone de montagne. La représentation spatiale de l'hétérogénéité à différentes échelles permet d'envisager de nombreuses applications dans les domaines de la conservation et de la gestion forestière multifonctionnelle. Nos résultats, relevant d'une approche spatialement explicite, valident l'hypothèse selon laquelle la qualité de l'habitat de la Gélinotte peut être modélisée avec une précision acceptable uniquement à partir d'indices paysagers, s'ils sont calculés sur la bonne étendue. Ils constituent donc une avancée pour une aide à la gestion, puisqu'ils préfigurent un outil permettant la localisation de zones prioritaires sur lesquelles la conservation de la Gélinotte (Tetrastes bonasia) peut être optimisée.
  • Article
    Purpose of ReviewUmbrella, flagship and keystone species are among the most widely employed surrogate species concepts. We explored whether these concepts are useful for understanding the consequences of landscape change. We assessed the literature on surrogate species in relation to landscape change and identified key foci and notable gaps within the existing evidence base. We outlined strengths and limitations of surrogate species as proxies for landscape change.Recent FindingsWe found that few studies evaluated whether taxa claimed to be surrogate species were in fact robust proxies. This is particularly so for flagship species but is also common in work on umbrella species. We also found marked differences in how the terms and concepts of umbrella, flagship and keystone species were used, both between studies and between disciplines (e.g. forestry versus community ecology). Research into surrogates is often conducted independently of research on landscape change. This leaves a major gap in knowledge about how surrogates can inform decision-making in relation to ongoing threatening processes, including landscape change.SummaryBased on results of our literature search and insights from large-scale, long-term empirical studies in south-eastern Australia, we identified a diverse mix of examples where the application of surrogate approaches has been successful, and where it has not. However, it is currently not possible to determine a priori where a given surrogate approach will work. Resolution of this problem requires considerable further work. Surrogates should be used in a critical way to help avoid mistakes in resource and biodiversity management.
  • Chapter
    The large felid carnivores are among the most endangered, and the most challenging, species to conserve on this increasingly human-dominated planet. In modern times, large felid carnivores were widely distributed in the continents of Asia, Africa, and the Americas. Unfortunately, global human expansion, loss of prey species, hunting and poaching, and retaliatory killings after livestock predation have greatly reduced and fragmented their original ranges and decimated their populations. In this chapter, large felid carnivore characteristics, usual habitats, ecology, and predatory behaviors are reviewed. Changes in great cat distribution, changes in wild prey populations resulting in a shift to increased livestock predation, and the resulting human-felid conflicts are discussed. Preservation of remaining wild large felid carnivore populations has become a global conservation priority as populations have plummeted over the last century. Current approaches to better understand and conserve these apex keystone predators and to maintain ecosystem integrity are discussed. Current strategies and policies to ameliorate and resolve the intricate and difficult problems of predator-human conflict are examined. The complex issues of “problem carnivores” and “man-eaters” are discussed. Finally, recommendations on creative, fluid, and scientifically sound strategies that might be employed to address these conflicts in a manner acceptable to all key stakeholders are discussed.
  • Article
    The minimal area of an animal populution is determined by A) the area requirement of reproductive units and B) by the viable population size. A) varies due to individually different and seasonally fluctuating home range sizes and is in addtiion strongly influenced by habitat quality. Population survival depends on deterministic as well as stochastic events and can therefore be estimated only with limited probability. A certain limitation of risk factors can be achieved by enlargement of the population size, increase in number of suitable habitats and reduction of isolation between inhabited areas. To determine the size of a “minimum viable population” (MVP) a “population vulnerability analysis” (PVA) is used as most important data base. The objectives of a MVP (e.g. 95% survival probability for the next 100 years) determines the necessary environmental conditions. A method which allows faster predictions was developed for special demands in practical implementation. A target species should be selected to give qualitative reasons for the protection of areas. Criteria for the selection of target species for conservation were developed which should be modified according to regional conditions. The concept of target species can also be used to quantify the evaluation of habitats scientifically, which is also an important step for management practises. The analysis of the data for selected species demonstrates the high variability of the area requirements, above all due to different habitat quality. For a MVP the area requirement is much higher than so far assumed (e.g. much more than 10 km2 for a passerine species). It is not possible to create a generally valid catalogue for the area requirements of species.
  • Chapter
    To maintain forest biodiversity, and if necessary to restore it, quantitative goals should be specified for all properties of a given landscape or region. Here an idea is presented on how biodiversity can be assessed in the boreal forest, which has a short history of transformation compared with other European forest types, and how quantitative nature conservation goals can be formulated in different geographical scales. Step 1 is to develop a book-keeping system for biodiversity. The starting point is the identification of the different disturbance regimes found in a natural boreal forest landscape. The links between site type and disturbance regime are critical because they shape the composition and structure of the forests, as well as many important processes, to which forest species have adapted. Step 2 is the development of practical methods to measure biodiversity in a landscape. The first task is to define the different ecological properties that can be found in a naturally dynamic boreal forest landscape as well as to identify species that are unique to each property. If it is possible to translate the species’ habitat requirements into criteria based on their environmental demands, these criteria can be used as opposed to the more costly inventories of certain indicator species in the field. It is important that any system and methods are validated in areas with original biodiversity. In step 3, when sufficient knowledge is available on how the amount of each property affects the viability of its indicator specie’s populations, strategic and operative goals can be formulated for each property representing the composition and structure, in different geographical scales, of naturally dynamic boreal landscapes. To implement this logic in practical management will alleviate the introduction of proactive management as well as monitoring of whether or not quantifiable criteria and indicator species change in the desired direction. The practical implementation is beyond the sphere of research but must be considered when developing methods to assess forest biodiversity.
  • Article
    Studies of the last few hundred years, at appropriate resolutions, are critical for understanding the dynamics of Australian ecosystems before and since European setlement in 1788 and for predicting effects of local, regional and global environmental changes over the next hundred years or so. -from Author
  • Article
    Most long-term studies of populations are now undertaken in remnants of what were once more extensive habitats. The unique characteristics of ecological systems on habitat islands must be considered in such studies. Those can include high edge to interior ratios, lacks of microhabitat heterogeneity, and other characteristics which can bias study results. Selective extinctions in habitat remnants may disrupt community interactions, effecting resource relationships, and the structures of surviving populations. Long-term studies of the population biology of Euphydryas editha in remnant native grasslands in California are presented as examples of the importance and difficulty of studying populations in habitat remnants. Studies on habitat remnants should be designed to enhance understanding of population processes leading to extinction and to aid in the design and management of reserves. -Authors
  • Article
    Full-text available
    Among the numerous site-ranking or reserve selection approaches is to select reserves based on the current distribution of a suite of species. This approach only requires distribution data for the species in the @'focal taxon.@' To what extent is the proposed reserve network dependent on the species included in the focal taxon? Can a suite of species act as an umbrella or indicator for other species whose distribution was not directly considered? A simple reserve selection algorithm was applied to species lists from sites where multiple taxa were sampled. The algorithm selected sites so that each species in the focal taxon was represented in at least one reserve. The species lists were complied for birds, mammals, reptiles, and plants on islands in the Gulf of California, and for birds, mammals, and plants in canyons in San Diego County, California. Within both data sets, there were significant rank correlations (P