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Abstract

The Andean Condor (Vultur gryphus) is a culturally iconic wildlife symbol for the South American Andes, but is naturally found at very low population densities, and is increasingly threatened. Using the Range Wide Priority Setting methodology, we (a group of 38 Andean Condor experts) updated the Andean Condor historical range (3,230,061 km2), systematized 9998 Andean Condor distribution points across the range, and identified geographic areas for which there was expert knowledge (66%), including areas where Andean Condors no longer occur (7%), and geographic areas where condors are believed to range, but for which there was not expert knowledge about condor presence (34%). To prioritize conservation action into the future and identify existing Andean Condor population strongholds, we used expert knowledge to identify 21 of the most important areas for the conservation of the species (i.e., Andean Condor Conservation Units [ACCUs]) that cover 37% of the revised historical range, and range in size from 837 km2 to 298,951 km2. In general, ACCUs were relatively small in the northern portion of the range in Venezuela, Colombia, Ecuador, and northern Peru, and significantly larger in the central and southern portion of the range in Peru, Bolivia, Chile, and Argentina, reflecting the reduced and narrower historical range in the northern portion of the range, as well as increased threats. Andean Condors can fly extremely long distances and so the populations of many neighboring ACCUs are probably still functionally connected, although this situation also underlines the need for integrated and large-scale conservation efforts for this species. As a function of the Range Wide Priority Setting results, we make recommendations to ensure population connectivity into the future and engage a wide range of actors in Andean Condor conservation efforts.
THE JOURNAL OF RAPTOR RESEARCH
A QUARTERLY PUBLICATION OF THE RAPTOR RESEARCH FOUNDATION, INC.
VOL.56 NO.1MONTH 2022
J. Raptor Res. 00(0):000–000
Ó2022 The Raptor Research Foundation, Inc.
DEFINING SPATIAL CONSERVATION PRIORITIES FOR THE ANDEAN
CONDOR (VULTUR GRYPHUS)
ROBERT B. WALLACE
1
,ARIEL REINAGA,AND NATALIA PILAND
Wildlife Conservation Society, Latin America and Caribbean Program, 2300 Southern Boulevard,
Bronx, NY 10460 USA
RENZO PIANA AND F. HERNA
´NVARGAS
The Peregrine Fund, 5668 West Flying Hawk Lane, Boise, ID 83709 USA
ROSA ELENA ZEGARRA
Servicio Nacional Forestal y de Fauna Silvestre (SERFOR-MINAGRI), Av. Javier Prado Oeste N82442, Urb.
Orrantia, Magdalena del Mar, Lima -17, Peru
SERGIO ALVARADO
Programa de Bioestad´ıstica, Instituto de Salud Poblacional, Facultad de Medicina, Universidad de Chile,
Santiago, Chile
and
Facultad de Ciencias de la Salud, Universidad de Tarapaca´ , Arica, Chile
SEBASTIA
´NKOHN
The Peregrine Fund, 5668 West Flying Hawk Lane, Boise, ID 83709 USA
and
Fundacio´n Co´ndor Andino, Tamayo N24-260 y Lizardo Garc´ıa, Quito, Ecuador
SERGIO A. LAMBERTUCCI AND PABLO ALARCO
´N
Grupo de Investigaciones en Biolog´
ıa de la Conservacio´n, INIBIOMA-CONICET, Universidad Nacional del
Comahue, Quintral 1250, Bariloche, Argentina
DIEGO ME
´NDEZ
Aves Rapaces en Bolivia—Programa de Investigacio´n, Calle El Villar 369, Sucre, Bolivia
and
Museo Nacional de Historia Natural, Calle 26 s/n, Cota Cota, La Paz, Bolivia
FAUSTO SA
´ENZ-JIME
´NEZ AND FRANCISCO CIRI
Fundacio´n Neotropical, Calle 2 #7—128 Cajica´, Cundinamarca, Colombia
JOSE
´´
ALVAREZ
MINAM, Direccio´n General de Diversidad Biolo´gica, Av. Antonio Miroquesada 425, Magdalena del Mar, Lima,
Peru
1
Emailaddress:rwallace@wcs.org
1
FERNANDO ANGULO
Divisio´n de Ornitolog´
ıa, Centro de Ornitolog´
ıa y Biodiversidad (CORBIDI), Santa Rita No. 105 Of. 202, Urb.
Huertos de San Antonio, Surco, Lima, Peru
VANESA ASTORE
Ecoparque Interactivo Buenos Aires, Rep ´
ublica de la India 3000, C1425FCF, Buenos Aires, Argentina
and
Fundacio´n Bioandina Argentina, Pasaje Juan de Castro 1457, CP 1406, Buenos Aires, Argentina
JANNET CISNEROS,JESSICA GA
´LVEZ-DURAND,AND ROSA VENTO
Servicio Nacional Forestal y de Fauna Silvestre (SERFOR-MINAGRI), Av. Javier Prado Oeste N82442, Urb.
Orrantia, Magdalena del Mar, Lima -17, Peru
CELESTE CO
´NDOR
CUNAMA, Jr. Chinchaysuyo 109, Departamento 201, Urb. Maranga San Miguel, Co´digo Postal 15088,
Lima, Peru
V´
ICTOR ESCOBAR
Corporacio´n Amigos del Co´ndor, Herna´n Corte´s 2953, dep.705, ˜
Nu˜
noa—Santiago, Chile, 7770074
MART´
IN FUNES,ALEJANDRO KUSCH,ADRIA
´NNAVEDA-RODR´
IGUEZ,CLAUDIA SILVA,
AND GALO ZAPATA-R´
IOS
Wildlife Conservation Society, Latin America and Caribbean Program, 2300 Southern Boulevard,
Bronx, NY 10460 USA
CAROLINA GARGIULO
Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Intendente G¨
uiraldes 2160—Ciudad
Universitaria—Pabello´n II, C.P.(C1428EGA), Capital Federal, Buenos Aires, Argentina
and
Centro de Zoolog´
ıa Aplicada, Universidad Nacional de Co´rdoba. Rondeau 798—Jard´
ın Zoolo´gico. C.P. (5000),
Co´rdoba, Argentina
SANDRA GORDILLO
Instituto de Antropolog´
ıa de Co´rdoba, Universidad Nacional de Co´rdoba—CONICET, Avenida Hipo´lito Yrigoyen
174, X5000 Co´rdoba, Argentina
JAVIER HEREDIA
Ecosistemas Argentinos, Calle 27 de Abril 2050 (5000), Co´rdoba, Argentina
RUBE
´NMORALES
Zoolo´gico de La Plata, Calle 52 y 118, Paseo del Bosque, Buenos Aires, Argentina
and
Fundacio´n Arreken, Calle 94 N 505 (esquina 5), La Plata, Buenos Aires, Argentina
ALEXANDER MORE
Naturaleza y Cultura Internacional, Urb. Laguna del Chipe, Mz C Lt 14, Piura, Peru
DAVID OEHLER
Nashville Zoo, 3777 Nolensville Pike, Nashville, TN 37211 USA
2 VOL. 00, NO.0
WALLACE ET AL.
OSCAR OSPINA-HERRERA
CORPOCALDAS, Calle 21 N823–22. Manizales, Caldas, Colombia
ANDRE
´SORTEGA
Universidad UTE, Facultad de Ciencias de la Salud & Carrera de Medicina Veterinaria,
Quito 170527, Ecuador
and
Fundacio´n Co´ndor Andino, Tamayo N24-260 y Lizardo Garc´
ıa, Quito, Ecuador
JOSE
´ANTONIO OTERO
El Huayco, Mza. o Lote. 11, Huachipa Norte, Lurigancho, Lima, Peru
CARLOS SILVA
Parque Zoolo´gico y Bota´nico Bararida, Gerencia de Manejo y Salud Animal, Av. Los Abogados con calle 13,
Barquisimeto, Venezuela
GUILLERMO WIEMEYER
Universidad de Buenos Aires, Av. Chorroar´
ın 280, C1427 CWN, Buenos Aires, Argentina
and
The Peregrine Fund, 5668 West Flying Hawk Lane, Boise, ID 83709 USA
LORENA ZURITA
Venancio Burgoa 978, La Paz, Bolivia
ABSTRACT.—The Andean Condor (Vultur gryphus) is a culturally iconic wildlife symbol for the South
American Andes, but is naturally found at very low population densities, and is increasingly threatened.
Using the Range Wide Priority Setting methodology, we (a group of 38 Andean Condor experts) updated the
Andean Condor historical range (3,230,061 km
2
), systematized 9998 Andean Condor distribution points
across the range, and identified geographic areas for which there was expert knowledge (66%), including
areas where Andean Condors no longer occur (7%), and geographic areas where condors are believed to
range, but for which there was not expert knowledge about condor presence (34%). To prioritize
conservation action into the future and identify existing Andean Condor population strongholds, we used
expert knowledge to identify 21 of the most important areas for the conservation of the species (i.e., Andean
Condor Conservation Units [ACCUs]) that cover 37% of the revised historical range, and range in size from
837 km
2
to 298,951 km
2
. In general, ACCUs were relatively small in the northern portion of the range in
Venezuela, Colombia, Ecuador, and northern Peru, and significantly larger in the central and southern
portion of the range in Peru, Bolivia, Chile, and Argentina, reflecting the reduced and narrower historical
range in the northern portion of the range, as well as increased threats. Andean Condors can fly extremely
long distances and so the populations of many neighboring ACCUs are probably still functionally connected,
although this situation also underlines the need for integrated and large-scale conservation efforts for this
species. As a function of the Range Wide Priority Setting results, we make recommendations to ensure
population connectivity into the future and engage a wide range of actors in Andean Condor conservation
efforts.
KEY WORDS:Andean Condor; Vultur gryphus; conservation;Range Wide Priority Setting.
DEFINICI ´
ON DE LAS PRIORIDADES ESPACIALES DE CONSERVACI ´
ON DE VULTUR GRYPHUS
RESUMEN.—El co´ ndor andino Vultur gryphus es un s´ımbolo de vida silvestre culturalmente ico´ nico para los
Andes sudamericanos, pero se encuentra naturalmente en densidades poblacionales muy bajas, y esta´ cada
vez ma´s amenazado. Utilizando la metodolog´ıa de Establecimiento de Prioridades de Conservacio´ n a lo largo
MONTH 0000 3
VULTURE SPATIAL CONSERVATION PRIORITIES
de su Distribucio´ n, nosotros (un grupo de 38 expertos en V. gryphus) actualizamos el a´rea de distribucio´n
histo´rica de la especie (3.230.061 km
2
), sistematizamos 9998 puntos de distribucio´ n en toda su a´rea de
distribucio´ n, e identificamos las a´reas geogra´ficas para las que hab´ıa conocimiento experto (66%),
incluyendo las a´reas en las que V. gryphus ya no esta´ presente (7%), y las a´reas geogra´ficas en las que se cree
que se distribuye, pero para las que no hab´ıa conocimiento experto sobre su presencia (34%). Para priorizar
las acciones de conservacio´ n en el futuro e identificar los bastiones poblacionales existentes de V. gryphus,
utilizamos el conocimiento experto para identificar 21 de las a´reas ma´s importantes para la conservacio´n de
la especie (es decir, Unidades de Conservacio´n del Co´ndor Andino [ACCU, por sus siglas en ingle´s]), que
cubren el 37% del a´rea de distribucio´n histo´rica, y tienen un tama˜
no de 837 km
2
a 298.951 km
2
. En general,
las ACCUs fueron relativamente peque˜
nas en la porcio´n norte del a´rea de distribucio´n en Venezuela,
Colombia, Ecuador y el norte de Per´
u, y significativamente ma´s grandes en la porcio´ n central y sur del a´rea
de distribucio´ n en Per´
u, Bolivia, Chile y Argentina, reflejando la reduccio´n y el estrechamiento del a´rea de
distribucio´ n histo´ rica en la porcio´ n norte de su area de distribucio´n,as´ı como el aumento de las amenazas. V.
gryphus puede volar distancias extremadamente largas, por lo que es probable que las poblaciones de muchas
ACCUs vecinas sigan estando conectadas funcionalmente, aunque esta situacio´ n tambie´n subraya la
necesidad de esfuerzos de conservacio´ n integrados y a gran escala para esta especie. En funcio´n de los
resultados del establecimiento de prioridades en toda el a´ rea de distribucio´ n, hacemos recomendaciones
para asegurar la conectividad de la poblacio´ n en el futuro y para involucrar a una amplia gama de actores en
los esfuerzos de conservacio´n de V. gryphus.
[Traduccio´ n de los autores editada]
INTRODUCTION
The Andean Condor (Vultur gryphus) is an emblem
of the Andean region and plays essential cultural
and ecological roles in Andean ecosystems. The
Andean Condor is found on both sides of the
Andean Mountain range, from Venezuela to Pata-
gonia (Birdlife International 2020). Condors are
apex scavengers that are able to open large carcasses
(Lambertucci et al. 2018), thereby speeding up
decomposition processes, and probably decreasing
risks of disease from carrion (Ogada et al. 2012).
Condors are among the longest-lived bird species
and are particularly vulnerable to extinction due to
their naturally low abundance, wide-ranging behav-
ior, late age of maturity, and low reproductive rate
(Lambertucci 2007, Houston et al. 2020).
Ongoing and persistent threats that impact
Andean Condor populations range from habitat
destruction, especially relevant for roosting and
nesting sites, to hunting due to perceived and
documented human-wildlife conflicts (Naller et al.
2008, Ministerio del Ambiente and The Peregrine
Fund 2018). Documented threats (Pavez and
Estades 2016) also include collisions with power line
infrastructure, an extremely worrying increase in
illegal carcass poisoning for predator control (Alar-
co´ n and Lambertucci 2018), secondary lead poison-
ing due to consumption of hunted prey species
(Wiemeyer et al. 2017), use of body parts in
handicrafts, collisions with vehicles (Speziale et al.
2008), competition for food from feral and domestic
dogs, tourism disturbance, and use in folkloric Yawar
Fiesta events in Peru, often resulting in serious injury
or death (Piana 2019).
The global Andean Condor population has been
proposed at 10,000 birds, including 6700 mature
individuals (Birdlife International 2020), although
there are no rigorous population estimates with
confidence limits across the majority of its distribu-
tion. Andean Condors have been virtually extirpated
from the northern portion of their range with fewer
than 300 adults now patchily distributed across
Ecuador, Colombia, and Venezuela (Sa´enz-Jime´nez
et al. 2014, Naveda-Rodr´ıguez et al. 2016, Vargas et
al. 2018), where reintroductions have been under-
way for more than a decade. In the central Andes,
recent efforts in Bolivia have estimated a total
population size of 1388 individuals (Me´ndez et al.
2015, 2019), and in a portion of Peru, between 155
and 249 individuals were estimated (Piana and
Angulo 2015). Population estimates of about 2000
individuals have been suggested for both Argentina
and Chile (World Wildlife Foundation and Funda-
cio´ n Bioandina 2000), although there are no
systematic counts in the field to support these
estimates, and given the geography of these neigh-
boring elongated countries, it seems probable that
there is overlap in the condor population between
countries. However, estimates at communal roosts in
Argentina and Chile suggest that there are hundreds
in each region studied (Feijo´ o 1999, Kusch 2004,
2006, Lambertucci 2010, Gargiulo 2014).
4 VOL. 00, NO.0
WALLACE ET AL.
National Red List classifications for the Andean
Condor are as follows: Critically Endangered in
Venezuela (Rodr´ıguez et al. 2015, Sharpe et al. 2015)
and Colombia (Renjifo et al. 2016); Endangered in
Ecuador (Freile et al. 2019), Peru (Decreto Supremo
004-2014-MINAGRI) and Argentina (Argentina Am-
biental 2017); and Vulnerable in Bolivia (Balderra-
ma et al. 2009) and Chile (Corporac´ıon Nacional
Forestal 1993). Given these national classifications
and the recent increase in the devastating threat of
illegal carcass poisoning for predator control, the
International Union for Conservation of Nature
(IUCN) recently upgraded the global red list
classification for the Andean Condor from Near
Threatened to Vulnerable (Birdlife International
2020).
There is an urgent need to identify priority
conservation areas for naturally scarce wildlife
species, especially those with large individual home
ranges, and low reproductive rates and genetic
variability (Santangeli et al. 2019). This is especially
important for species under known and significant
threats and that are particularly sensitive to environ-
mental change due to their longevity (Beissinger
2000, Owens and Bennett 2000, Knight and Cowling
2007). Andean Condors are known to use large areas
covering a variety of management jurisdictions;
studies in Argentina have demonstrated the impor-
tance of protected areas in the protection of the
species, and some researchers have proposed prior-
itization of conservation areas based on movement
data of satellite-tagged individuals (Lambertucci et
al. 2014, Guido et al. 2020, Perrig et al. 2020).
However, there is a pressing need for a prioritization
of conservation areas across the entire distribution
of the species.
The main objective of this work was to inform
future multi-national conservation decision-making
by systematizing existing distributional knowledge
and identifying priority conservation areas for the
Andean Condor throughout its distribution. To do
so, we selected the expert-driven Range Wide
Priority Setting exercise (RWPS), which was devel-
oped to systematize scarce and usually dispersed data
regarding the overall distribution of globally threat-
ened wildlife species, to make informed manage-
ment decisions regarding their conservation
(Sanderson et al. 2002, Thorbjarnarson et al. 2006,
Marieb 2007, Taber et al. 2009, Wallace et al. 2014b).
The result of the RWPS was the report Saving the
Symbol of the Andes: A Range Wide Conservation Priority
Setting Exercise for the Andean Condor (Vultur gryphus;
Wallace et al. 2020). This manuscript is a summary of
that report.
METHODS
In 2015, a partnership of the National Forestry and
Wildlife Service of Peru, the Peruvian Ministry of
Environment, The Peregrine Fund, and the Wildlife
Conservation Society launched an Andean Condor
Range Wide Priority Setting exercise with 38 experts
working with the species in different parts of its
distribution.
Prior to an in-person May 2015 workshop, we
systematized information about Andean Condor
distribution from three sources. Firstly, we collected
Andean Condor distribution points derived from a
thorough literature review. To do so we gathered
information by using key words (Andean Condor,
co´ ndor andino, Vultur,Vultur gryphus) to search
GoogleScholare, as well as unpublished reports
from Bolivia and Peru, which had been gathered in a
previous systematization effort for the central
portion of the range (Wallace et al. 2014a).
Secondly, we requested all Andean Condor occur-
rence records from the largest citizen science effort
in the world, eBird (https://ebird.org/explore),
and downloaded them in May 2015. Finally, we
solicited additional unpublished information from
Andean Condor experts from different countries
using a specific spreadsheet on observed Andean
Condor roosts, nests, and feeding sites (Supplemen-
tal Material). By comparing the location, date and
observer information for distribution points, we
eliminated duplicate distribution points arising
from more than one source. In the mapping analyses
described below, we used all unique distribution
points. All current Andean Condor distribution
areas were also assumed to be included in the
historical distribution.
As part of the RWPS methodology (Sanderson et
al. 2002), prior to the workshop Andean Condor
experts were also asked to draw polygons describing
their knowledge of Andean Condors including: (1)
Area of Knowledge (areas where experts are
knowledgeable enough to express opinion about
the presence or absence of Andean Condors); (2)
Proposed Actual Distribution (areas where experts
believe the Andean Condor has occurred in the last
20 yr), and (3) Andean Condor Conservation Units
(ACCUs; areas perceived by experts to be important
strongholds for the long-term conservation of
Andean Condor populations within the expert’s
area of knowledge). Experts received maps of their
MONTH 0000 5
VULTURE SPATIAL CONSERVATION PRIORITIES
country in GoogleEartheformat as an additional
tool with which to draw polygons and/or place
distribution points. We processed and combined all
information into one Geographic Information
System (GIS) using the ArcGIS platform (Version
10.3) and incorporated data on human settlements;
international, state and provincial boundaries; and
main and secondary roads (DIVA-GIS; https://www.
diva-gis.org/Data), as well as satellite images to
facilitate recognition of physical characteristics such
as mountains, valleys, and plains.
Workshop participants (n¼101) were placed into
seven geographical working groups: (1) Colombia
and Venezuela, (2) Ecuador, (3) northern Peru, (4)
southern Peru, (5) Bolivia, (6) Argentina, and (7)
Chile. Using printed maps and satellite imagery
(http://www.esri.com/software/arcgis/arcgisonline/
bing-maps.html) at varying scales depending on the
country (from 1:900,000 for Bolivia to 1:5,500,000
for Chile), and digital versions of the same on
portable computers, each group was asked to review
the historical range of the Andean Condor (Fjelds˚a
and Krabbe 1999). Experts used their collective
knowledge about Andean Condors in combination
with all systematized distribution points, altitudinal
contours for the range, and satellite imagery to
define the historical range (where Andean Condors
are thought to have occurred since 1900).
The working groups were then asked to review pre-
workshop draft maps based on information provided
by experts prior to the workshop on the conservation
status of the Andean Condor. First, working groups
reviewed systematized distribution data divided into
historical (1900–1994) and current distribution
points (1995–2015). Then the working groups
defined places or polygons where experts had
knowledge of the condor, ranging from isolated
observations, to information on movement data
from an individual condor, to the presence of a nest
or communal roost, to in-depth ecological knowl-
edge, and distinguished places within the historical
range where Andean Condors are known to no
longer occur. Working groups also identified places
where experts did not have knowledge of either
presence or absence of Andean Condors. Finally,
working groups proposed ACCUs within their area
of knowledge. Groups worked in a systematic order
and clearly marked changes on printed satellite
image maps and/or digital versions in kmz format
(Google Earthe). Neighboring working groups
then met to compare, discuss and integrate distri-
bution, knowledge, and ACCU polygons for a
number of transboundary areas across the range.
Subsequently, maps for the seven working groups
were combined into range-wide maps for each
category (historical distribution, expert knowledge,
no longer occur, without expert knowledge, and
ACCUs).
After the workshop, the maps were digitized and
modified according to the corrections and proposals
of the workshop participants. We then sent the
modified maps to the participating Andean Condor
experts for review and incorporated any final edits.
We divided the ACCUs into three size classes: (1)
relatively small ACCUs of ,20,000 km
2
, (2) medium-
sized ACCUs of 20,001 to ,100,000 km
2
, and (3)
relatively large ACCUs of .100,000 km
2
. We also
calculated the percentage of each ACCU protected
by three jurisdictional categories of protected areas:
(1) National Protected Areas, (2) State or Regional
Scale Protected Areas, and (3) Municipal or Private
Protected Areas, by overlapping the ACCUs and
protected areas using the clip and intersect tools in
ArcGIS.
RESULTS
Historical Range of the Andean Condor. Overall,
the revision of the Andean Condor historical range
resulted in a polygon of 3,230,061 km
2
(Fig. 1a).
This represents an almost 37% increase for the
estimated historical range as compared to the
previously estimated area of approximately
2,362,397 km
2
(Fjelds˚a and Krabbe 1990). From this
point forward, ‘‘historical range’’ is used to refer only
to the 2020 revised historical range map.
Andean Condor Distribution Points. The litera-
ture review resulted in a total of 928 distribution
points as a baseline prior to the workshop. Andean
Condor experts attending the RWPS exercise
provided an additional 793 distribution points to
the database. After filtering duplicate points from
the database, eBird added 8277 distribution points.
The overall database of Andean Condor distribution
points consisted of 9998 data points (historical
1900–1994: 185 data points; current 1995–2015:
9813 data points). The number of Andean Condor
distribution points for each country ranged from 12
distribution points in Venezuela, the northern
distributional extreme, to 3863 data points in Chile
at the southern extreme (Table 1).
The portion of the species’ overall historical range
found within each country ranged from ,1% in
Venezuela to almost 43% in Argentina. The south-
ern portion of the historical range in Argentina and
6 VOL. 00, NO.0
WALLACE ET AL.
Figure 1. Revised Andean Condor historical range (a) as compared to Fjelds˚a and Krabbe (1999) previous historical
range, and (b) highlighted areas with and without expert knowledge for Andean Condors, and areas where they no longer
occur. Figure adapted from Wallace et al. 2020.
MONTH 0000 7
VULTURE SPATIAL CONSERVATION PRIORITIES
Chile accounted for just over 66% of the overall
revised historical range. The central portion in Peru
and Bolivia represented almost 28%, and the
northern portion of the range in Colombia, Ecua-
dor, and Venezuela accounted for just 6% of the
historical range (Table 1). The map of distribution
points (Fig. 2) reveals a virtually continuous distri-
bution for the Andean Condor in Argentina, Chile,
Bolivia, Peru, and Ecuador, with the distribution
points then being scarcer and more isolated in
Colombia and Venezuela.
Andean Condor Extirpated Areas. Workshop
participants identified areas where Andean Condors
no longer occur within their historical range (Fig.
1b), amounting to just over 7% of the estimated
historical range. These areas consisted of two small
polygons in the central Andes of Colombia, a similar-
sized polygon in the central portion of the Ecuador-
ian Andes, three larger polygons in northern Peru
from the Pacific to the eastern Andes, one polygon
in the Sajama region of Bolivia, and the two largest
polygons in southeastern Argentina along the
Atlantic coast.
Areas Identified With and Without Andean Condor
Expert Knowledge. Andean Condor experts detailed
areas with or without expert knowledge (Fig. 1b).
Unsurprisingly, the polygon areas with knowledge
across the Andean Condor distribution largely
reflect known localities (Fig. 2). Overall, experts
identified almost 66% of the Andean Condor
historical range as either containing Andean Con-
dor populations (59%), or where Andean Condors
are now considered absent (7%; Table 2). Experts
noted that 34% of the historical range is without
expert knowledge (Table 2). Nevertheless, the eBird
dataset provided presence data for large portions in
the southern part of the range (Argentina and
Chile), in areas identified as without expert knowl-
edge or expert-derived distribution points (Fig. 2).
Priority Andean Condor Conservation Units (AC-
CUs). In total, in the original workshop 31 ACCUs
were proposed by Andean Condor experts, covering
an area representing 37% of the adjusted historical
range. More than half of the area prioritized as
ACCUs are in Argentina (51%), slightly more than
would be expected based on the portion of the
historical range for Argentina (43%; Table 1).
Experts also prioritized a larger area in Bolivia
(18%) than might be expected based on the
percentage of the historical range (11%), whereas
in Peru (14%), and particularly Chile (11%), experts
prioritized less area than might be expected given
the historical range in those countries, 16% and 23%
respectively (Tables 1, 3). ACCUs in the northern
part of the Andean Condor range (Venezuela,
Colombia, and Ecuador) were small (just under
6%) and thus reflect small historical range portions
(just over 6%).
Several ACCUs in neighboring countries were
immediately adjacent to each other; in the post-
workshop analysis we combined several of the
original country-specific ACCUs into trans-boundary
ACCUs. Once these were combined, the number of
ACCUs was reduced to 21 units (Table 4; Fig. 3).
Fourteen of the ACCUs (most in Venezuela, Colom-
bia, Ecuador, and northern Peru) were classified as
small (Table 4), three as medium, and four as large
(three of which are enormous transboundary areas
Table 1. Andean Condor revised historical range area and
distribution points by country (Wallace et al. 2020).
COUNTRY
REVISED HISTORICAL
ANDEAN CONDOR
RANGE SIZE
(km
2
)
PERCENT OF
REVISED
HISTORICAL
RANGE
NUMBER OF
DISTRIBUTION
POINTS USED
IN RWPS
Venezuela 17,656 0.55 12
Colombia 129,867 4.02 222
Ecuador 53,831 1.67 1163
Peru 529,097 16.38 939
Bolivia 366,091 11.33 1181
Chile 751,481 23.27 3863
Argentina 1,382,037 42.78 2639
Total 3,230,061 100 9998
Table 2. Andean Condor expert knowledge across the revised historical range (Wallace et al. 2020).
POLYGON TOTAL AREA (km
2
)
PERCENT OF
REVISED HISTORICAL RANGE
Area with expert knowledge where Andean Condors still occur 1,888,924 58.5
Area with expert knowledge where Andean Condors no longer exist 236,012 7.3
Area without expert knowledge 1,105,125 34.2
Total revised historical Andean Condor range 3,230,061 100.0
8 VOL. 00, NO.0
WALLACE ET AL.
Figure 2. Distribution of confirmed Andean Condor localities obtained from: (a) Andean Condor experts during the
Range Wide Priority Setting (RWPS) exercise; (b) a literature review prior to the RWPS; (c) known nesting and roosting
sites; and (d) eBird occurrences up to May 2016. Figure adapted from Wallace et al. 2020.
MONTH 0000 9
VULTURE SPATIAL CONSERVATION PRIORITIES
of well over 200,000 km
2
, one bridging Argentina
and Bolivia and two running along the Argentina
and Chile border). A total of nearly 16% of the area
defined as ACCUs is under formal protection,
although there is considerable variation in protec-
tion among ACCUs (Table 4), protected percentages
varying between 0% to .50% for smaller ACCUs,but
mainly well below 20% protection for the medium
and large ACCUs (range ¼9–30%). Nevertheless, at
least 30% of the area of seven of the ACCUs is already
protected, although that includes only one of the
seven largest ACCUs (Table 4).
DISCUSSION
Andean Condor Historical Range. The historical
range update included a major increase in area and
the number of observations available, primarily due
to records from eBird, and the fact that the spatial
information used for this exercise was of consider-
ably better quality and finer scale because of the
advent of GIS technology and increased availability
of satellite images. This updated species global
distributional range totals 3,230,061 km
2
and repre-
sents an important perspective with which to set
future conservation targets, as well as measure
changes in the species’ distribution. The Andean
Condor has an extremely linear distribution along
the Andes (Fjelds˚a and Krabbe 1999, Houston et al.
2020), which might make it more prone to range
fragmentation and loss of gene flow.
Expert Knowledge Coverage within the Andean
Condor Historical Range. The Andean Condor
experts felt comfortable expressing their opinion
about Andean Condor presence in almost 59% of
the revised historical range and absence in 7% of the
revised historical range. This amounted to a total
knowledge coverage of 66%, or almost two-thirds of
Table 3. Size and area percentage of Andean Condor
Conservation Units (ACCUs) by country (Wallace et al.
2020).
COUNTRY n
AREA (km
2
)
IN ACCUS
%TOTAL
CONSERVATION
UNIT AREA
Venezuela 2 12,447 1.03
Colombia 8 31,492 2.62
Ecuador 3 22,973 1.91
Peru 5 169,131 14.05
Bolivia 3 213,698 17.75
Chile 5 138,058 11.47
Argentina 6 615,903 51.17
Total area in ACCUs 31 1,203,703 100
Table 4. Final list of Andean Condor Conservation Units (ACCUs; Wallace et al. 2020).
ACCU# COUNTRY ACCU NAME
AREA
(km
2
)
TOTAL %
PROTECTED
1 Venezuela Cordillera de Me´rida 11,870 39.0
2 Venezuela and Colombia Serran´ıa del Perija´ 3068 17.7
3 Colombia Sierra Nevada de Santa Marta 9289 55.9
4 Colombia Pa´ramos de los Andes Nororientales 5593 32.1
5 Colombia Corredor de Pa´ramos Guantiva-La Rusia-Iguaque 837 7.7
6 Colombia Chingaza 1381 57.8
7 Colombia Los Nevados 8542 9.5
8 Colombia Purace´ 2487 46.6
9 Colombia and Ecuador Chiles-Llanganates 13,349 33.6
10 Ecuador Sangay National Park 3220 55.6
11 Ecuador Azuay-Loja-El Oro 7278 3.7
12 Peru Andes de Piura-Lambayeque 8125 0
13 Peru Illescas 1813 0
14 Peru Andes Centrales de Per´
u 66,131 9.2
15 Peru and Bolivia Sur de Per ´
u-Norte de Bolivia 97,017 10.9
16 Bolivia ıpez-Sillillica 17,538 30.6
17 Argentina and Bolivia Cordillera Oriental-Sierras Subandinas-Sierras Pampeanas 298,951 16.5
18 Argentina and Chile Andes Centrales-Sierras Pampeanas 215,079 12.1
19 Argentina and Chile Patagonia Norte 237,784 12.9
20 Argentina Somuncura´ 53,187 30.1
21 Argentina and Chile Patagonia Sur 141,167 15.9
10 VOL. 00, NO.0
WALLACE ET AL.
Figure 3. Priority Andean Condor Conservation Units (UCCA in Spanish) across the range in South America. Figure
adapted from Wallace et al. 2020.
MONTH 0000 11
VULTURE SPATIAL CONSERVATION PRIORITIES
the historical range. Nevertheless, in all countries,
there were significant areas without expert knowl-
edge about Andean Condors, totaling 34% of the
historical range (notably in Bolivia, Peru, and
Chile). In addition, many areas with expert knowl-
edge were limited to Andean Condor observations
and lacked detailed ecological or population infor-
mation.
Expert knowledge coverage of 66% is lower than
other iconic species previously considered in com-
plete RWPS exercises in the region. For example, the
original jaguar (Panthera onca) RWPS analysis re-
vealed expert knowledge areas covered 83% of the
historical range (Sanderson et al. 2002), which
increased in 2006 to 96% (Marieb 2007). For less
cryptic species, expert knowledge covered 99% of
the historical range for white-lipped peccaries
(Tayassu pecari) and almost 100% for lowland tapirs
(Tapirus terrestris; Taber et al. 2009). However, expert
knowledge coverage was just 58% for Andean bears
(Tremarctos ornatus) in Bolivia and Peru (Wallace et
al. 2014b), which also have an exceptionally linear
distribution that is largely confined to the eastern
slopes of the Andes Mountain range from Venezuela
to Bolivia.
In the northern and central portions of the range
there was considerable overlap and agreement
between expert-driven knowledge and available
eBird data. However, in the southern portion of
the range, especially in Chile, large areas were
identified as areas without expert knowledge, but
were populated with significant concentrations of
Andean Condor observations from eBird (Fig. 2d).
Future studies in Chile should verify locations with
high densities of citizen science observations.
Andean Condor Actual Range. Workshop partici-
pants identified eight polygons where Andean
Condors are considered to have been extirpated:
two in central Colombia, one in south-central
Ecuador, two in northern Peru, one in western
Bolivia, and two in southeastern Argentina, account-
ing for 7% of the revised historical range. Given that
34% of the revised historical range has no knowl-
edge coverage from participating experts, the need
for further expert participation and/or fieldwork is
evident and pressing. Thus, current knowledge
suggests that Andean Condors remain present in at
least 59% of their historical range, although this
increases to around 80% when incorporating eBird
knowledge (Fig. 2d). Serious threats, such as large
poisoning events with pesticides (Alarco´n and
Lambertucci 2018, Estrada Pacheco et al. 2020)
and lead (Wiemeyer et al. 2017), in combination
with confirmed local extirpations herein, under-
score the need for species-specific conservation
planning and actions.
In comparison to tapirs, peccaries, and jaguars in
South America, all of which have been extirpated
from between 14 and 39% of their historical range
(Marieb 2007, Taber et al. 2009), the 7% value for
condors is encouraging. However, in this work we
only evaluated the presence of individuals, and we
do not know their regional abundances, their
demographic composition or ability to breed, or
their ability to perform historical behaviors, such as
foraging movements (Lambertucci et al. 2012,
2018). Andean Condors have overlapping individual
home ranges, which are also orders of magnitude
larger (Lambertucci et al. 2014) than those of tapirs
(Medici 2011), white-lipped peccaries (Moreira-
Ram´ırez et al. 2019), or jaguars (Morato et al.
2016), and many condors can be vulnerable to
threats in a relatively small area. Therefore, the
percentage of historical range where Andean Con-
dors are still observed as a standalone indicator of
the conservation status of this species may not be
appropriate.
Andean Condor Conservation Units (ACCUs).
Andean Condor experts proposed a total of twenty-
one ACCUs from western Venezuela to Tierra del
Fuego, which may represent the best hope for the
long-term conservation of Andean Condors across
the actual range. The ACCUs cover 37% of the
estimated actual range of the species. Experts
defined ACCUs ranging from relatively small areas
of just 837 km
2
(Corredor de Pa´ramos Guantiva-La
Rusia-Iguaque) to massive areas of up to 298,951
km
2
(Cordillera Oriental-Sierras Subandinas-Sierras
Pampeanas; Table 4). These immense areas high-
light the need for an integrated and landscape-level
approach for Andean Condor conservation involv-
ing a diversity of jurisdictions and associated local
stakeholders.
In general, ACCUs are relatively small in the
northern portion of the Andean Condor range
(Venezuela, Colombia, Ecuador, and northern
Peru), and an order of magnitude larger in the
central and southern portion of the range (central
and southern Peru, Bolivia, Chile, and Argentina),
which to a degree reflects the pattern of the
historical distribution of the species. Several of the
smaller ACCUs are almost certainly not large
enough to permanently hold viable populations of
Andean Condors, given the birds’ exceptionally
12 VOL. 00, NO.0
WALLACE ET AL.
large ranging patterns (Lambertucci et al. 2014).
However, within the framework of a regional analysis
for a wide-ranging species, these sites are important
as they have known roosting, nesting, and feeding
sites (Lambertucci 2010, Perrig et al. 2020). Thus,
recognition of these critical sites in the northern
portion of the range, where Andean Condors are
especially threatened, is an important step forward
in conservation planning for the species. Designat-
ing movement corridors as priority areas for
conservation may be less important for Andean
Condors than terrestrial animals due to their
capacity for rapid travel across huge home ranges
(Lambertucci et al. 2014) and resulting low genetic
variability (Hendrickson et al. 2003, Padro et al.
2018).
The medium and large ACCUs in the central and
southern portions of the historical range may well be
large enough to permanently hold meaningful
populations of Andean Condors. However, the
ACCUs’ immense sizes underline the need for
integrated conservation approaches that embrace
the importance of including different types of
protected areas (e.g., international biosphere re-
serves; Guido et al. 2020) and also going beyond
protected area limits and engaging in discussions
and management practices with a wide range of local
stakeholders and communities. It is also worth
stressing that given the flying capacity of Andean
Condors, populations of many neighboring ACCUs
are probably still functionally connected. Nonethe-
less, establishing longer-term connectivity through
strategic management activities should be consid-
ered, especially for the smaller ACCUs.
Currently, 14% of the Andean Condor’s historical
range is under formal protection. This does not
meet the 17% recommended by the Convention on
Biological Diversity as a 2011–2020 goal in the Aichi
targets (Convention on Biological Diversity 2011). It
is important to emphasize that individual Andean
Condors travel huge distances, and the ranges of
most individuals in the global population include
protected areas, but also large portions of other
unprotected types of land management (Lamber-
tucci et al. 2014, Guido et al. 2020). The protected
portions of the ACCUs were deemed to be especially
important for nesting and roosting. However, given
that adult mortality is thought to be the most
important driver of population rate change in
Andean Condors, on-the-ground conservation ef-
forts should focus on changing human behavior in
condor foraging areas, which concentrate many of
the most important threats such as poisoning
(Lambertucci et al. 2014, Naveda-Rodr´ıguez et al.
2016, Guido et al. 2020, Perrig et al. 2020). Thus, as
for Old World vultures (Santangeli et al. 2019), the
challenge into the future will be to secure the
sustainable and effective management of the pro-
tected areas, as well as to effectively achieve
conservation outside of protected areas within the
ACCUs.
In summary, the results demonstrate a clear
pattern reflected across the historical range, current
range, as well as identified ACCUs. The areas from
the northern portion of the historical range are
significantly smaller and substantially more frag-
mented than in the central range, and especially the
southern portion of the range (Fig. 3). This pattern
is also reflected in terms of known data regarding
Andean Condor populations, with numbers in
Venezuela, Colombia, and Ecuador particularly low
as compared to Peru, Bolivia, Chile, and Argentina.
The RWPS exercise amassed almost 10,000 distribu-
tion points and this data set, along with the
associated polygons, provides a means of monitoring
the conservation status of Andean Condors across
the range into the future. Indeed, as satellite
telemetry studies increase, the possibility of incor-
porating an even richer data set regarding condor
flight paths, roosts, nests, and foraging sites into
conservation decision-making processes (Perrig et
al. 2020) will allow further refinement of these
analyses.
Next Steps and Recommendations. Based on the
results of this Range Wide Priority Setting exercise
for the Andean Condor, we propose the following
priority recommendations: (1) develop specific and
comprehensive analyses and conservation plans with
integrated and diverse conservation actions for
identified ACCUs; (2) conduct surveys regarding
the presence of Andean Condors in areas without
expert knowledge about the species, or with very
poor knowledge within existing ACCUs; (3) design
and apply standardized Andean Condor census
methodologies using examples from Bolivia, Chile,
Ecuador, Argentina, and Peru, but also other
monitoring techniques (e.g., Perrig et al. 2019).
These standardized methodologies should then be
applied across the range, to assess population size,
especially at priority conservation sites, thereby
better informing future conservation decision mak-
ing processes; (4) encourage greater international
collaboration and interaction between countries, as
Andean Condors do not recognize borders and
MONTH 0000 13
VULTURE SPATIAL CONSERVATION PRIORITIES
require conservation across various jurisdictions; (5)
promote mixed conservation strategies that recog-
nize the role that local communities and private
landowners will play in overall Andean conservation
across the entire distribution area, and the need to
increase environmental education and outreach; (6)
work with the governments of the Andean nations to
address the most pressing threats to Andean Condor
populations, especially including legislation to pros-
ecute the use of poisons in carcasses, and eliminate
the risks of lead poisoning.
SUPPLEMENTAL MATERIAL (available online):
Spreadsheet S1: Localidades, Amenazas y Unidades
Prioritarias para la Conservacio´n del Co´ndor Andino.
ACKNOWLEDGMENTS
We thank all participants at the II International
Symposium on the Andean Condor held in Lima, Peru,
in May 2015. We are especially grateful for the generous
financial support provided by the National Forestry and
Wildlife Service of the Ministry of Agriculture in Peru, and
the Ministry of the Environment in Peru, as well as
significant matching funds from the Wildlife Conservation
Society and The Peregrine Fund. Avianca Airlines kindly
supported the airfares of several of the international
condor experts, ensuring a workshop with adequate
participation of experts from across the region. We thank
Evan Buechley, Todd Katzner, and one anonymous
reviewer for comments and suggestions that greatly
improved an earlier version of this report.
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16 VOL. 00, NO.0
WALLACE ET AL.
... The Andean condor is a large obligate scavenger that inhabits the Andes Mountain range in South America (from Venezuela to the south of Chile and Argentina) [34,41]. Condor populations have suffered a retraction in some areas, and their populations are tending to decrease [33,34,42]. Their reproductive strategy is one of the slowest among birds; they commonly lay one egg every two years, then the chick spends up to six months in the nest, and stays with its parents until 15 months of age [27,43]. ...
... This highly valuable information is necessary for a better understanding of the movement ecology of the species, and its association with, for instance, anthropogenic threats [33]. It is also relevant to the design of management strategies and conservation action [35,42,80]. The information provided can also be useful for other species that cross national and international borders, given that conservation action and management policies often differ between countries [60,81]. ...
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... Due to anthropogenic threats such as habitat degradation, poisoning with pesticides, lead intoxication, illegal capture, free-ranging dogs and shooting [21,[82][83][84], Andean Condors and California Condors, listed as Vulnerable and Critically Endangered [23], respectively, have suffered considerable population declines and geographic range contractions [85][86][87][88]. Furthermore, it is also expected that climate change will cause Black Vultures to move to higher altitudes and this will consequently increase overlap and competition with Andean Condors [89]. ...
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The New World Vultures (Cathartidae) include seven species of obligate scavengers that, despite their ecological relevance, present critical information gaps around their evolutionary history and conservation. Insights into their phylogenetic relationships in recent years has enabled the addressing of such information gaps through approaches based on phylogeny. We reconstructed the ancestral area in America of the current species using two regionalization schemes and methods: Biogeography with Bayesian Evolutionary Analysis (BioGeoBears) and Bayesian Binary Model–Monte Carlo Markov Chains (BBM–MCMC). Then, we identified the priority species and areas for conservation by means of the Evolutionary Distinctiveness index (ED), as a proxy of the uniqueness of species according to phylogeny, and the Global Endangerment index (GE), mapping phylogenetic diversity. We found that the ancestral area of NewWorld Vultures in America corresponds to South America, with dispersal processes that led to a recolonization of North America by Coragyps atratus, Gymnogyps californianus and Cathartes aura. We identified the Black Vulture, G. californianus and Vultur gryphus as priority species based on ED and “Evolutionary Distinct Globally Endangered” (EDGE) indexes, and the lowlands of Amazon River basin and the Orinoco basin and some tributaries areas of the Guiana Shield were identified as the priority areas when mapping the phylogenetic diversity. This study highlights the importance of filling knowledge gaps of species of conservation concern through the integration of evolutionary and ecological information and tools and, thus, developing adequate strategies to enhance the preservation of these species in the face of the current loss of biodiversity.
... In addition, we used the resistance surface map to model a conservation corridor across the northern Andes, from the potential source population of Peru to the small populations in Ecuador and Colombia. We calculated the probability of passage between major roosting sites in the region (Piana and Vargas, 2018;Wallace et al., 2022) using random-walk paths estimated in the gdistance R-package ( van Etten, 2017). Finally, we calculated the percentage of protected areas and land cover use type (e.g., grasslands, woods, urban zones) intersected by the resistance paths throughout the identified corridor (Table S1). ...
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