Project BioMap: documenting the global museum resource of Colombian

birds for research and conservation

Proyecto BioMap: documentando los recursos de museos de aves colombianas para investigación y

conservación

Juan–Carlos Verhelst–Montenegro1,2*, Paul Salaman3** and Robert Prys–Jones4

1
Conservation International – Colombia. Cra. 13 No. 70–41, Bogota
2
Geography Department, King’s College London. Strand, London WC2R 2LS
3
Conservation International. 2501 M Street NW, Suite 200, Washington DC 20037, USA
4
Bird Division, The Natural History Museum, Akeman St, Tring, Herts HP23 6AP, UK

Current addresses
*
Atlas of the Birds of Colombia https://sites.google.com/site/haariehbamidbar/
**
Rainforest Trust 7078 Airlie Rd., Warrenton, VA 20187, USA

Email: juan.verhelst@gmail.com

Resumen — El Proyecto BioMap fue una iniciativa transnacional entre instituciones

del Reino Unido (RU), Colombia y los Estados Unidos de América (EUA), que dio
inicio en 2001. El objetivo de dicho proyecto fue digitalizar y verificar todos los
especímenes de aves colombianas depositados en museos de historia natural alrededor
del mundo. Información de un total de 217 659 especímenes de aves fue compilada en
una base de datos georreferenciados (siempre que fue posible), la cual fue publicada en
el Internet (http://biomap.net). Los colectores más importantes fueron M. A. Carriker
Jr. y K. Von Sneidern, habiendo colectado alrededor de 28 000 especímenes cada uno.
Los períodos más activos de colecta fueron entre los años 1910–1920 y 1940–1980. En
el primer período destacándose las expediciones realizadas por F. M. Chapman para el
AMNH, mientras que durante el segundo los varios esfuerzos liderados por
investigadores e instituciones nacionales. Análisis preliminares de la información
mostraron que cerca del 55% de los especímenes colombianos están en los EUA, la
gran mayoría en solo cinco museos, 33% en Colombia y solamente un 12% en museos
del RU y Europa. Un total de 1815 especies están representadas en colecciones, un
95% del total de las especies registradas en el país. Además, 9705 y 2855 especímenes
corresponden a 180 especies de rango restringido y a 102 especies globalmente
amenazadas (categorías CR, EN y VU de la UICN). Por otro lado, para

Notulae Ornitologicae Columbianae 2

 Project BioMap – Colombia

aproximadamente 80% de las localidades de colecta hay menos de 50 especímenes,

señalando una baja tasa de muestreo al nivel puntual de localidad. El análisis de la
densidad del número de especímenes colectados en cuadrículas demostró que a un
tamaño de cuadrícula de ≈15 km cerca del 80% del país no ha tenido ninguna actividad
de colecta y que para solo el 1% del territorio colombiano se han colectado más de 500
especímenes, mientras que usando un tamaño de cuadricula de ≈60 km solamente para
el 11% del país se han colectado más de 1000 especímenes. De manera contraria a lo
esperado, recientemente para dos de las cuatro localidades con mayor número de
colectas (>3000 especímenes) en Colombia se han descrito nuevas especies para la
ciencia, las cuales no habían sido colectadas previamente. Estimamos que posiblemente
más de 100 taxa de aves nuevos para la ciencia pueden estar aun esperando ser
descubiertos y descritos en Colombia. Considerando las amenazas a las especies de
rango restringido, es urgente continuar en Colombia los inventarios de aves. Así
mismo, como realizar colectas de especímenes en el campo, pero de forma
extremadamente cuidadosa, dirigida y ética. También es importante mantener,
fortalecer y coordinar las actividades de colecta de aves de los diferentes museos
activos en el país. El establecimiento de la base de datos Darwin–Hernández por parte
del Proyecto BioMap ha creado una síntesis de la información existente de los
especímenes de aves colombianos, la cual está disponible en el Internet para todas las
personas e instituciones interesadas en priorizar futuros esfuerzos de investigación y
acciones de conservación en Colombia.

Palabras clave — BioMap; alianza internacional; biodiversidad; aves; colecciones;

Colombia

Abstract — Project BioMap, a tri–national initiative between British (UK), Colombian

and United States of America (USA) institutions, began in late 2001. The project aim
was to digitise and verify all Colombian bird specimens deposited in natural history
museums around the world. A total of 217 659 Colombian bird specimens in 87
museums were databased and georeferenced (whenever possible) and made available
online (http://biomap.net). The most important collectors were M. A. Carriker Jr. and
K. Von Sneidern, with about 28 000 specimens each. The most active periods of
collection were from 1910–1920 and 1940–1980, the former period marked by
expeditions led by F. M. Chapman for the AMNH and the latter by collecting efforts
led by national researchers for Colombian museums. Preliminary analyses of the
information showed that about 55% of Colombian specimens are held in the USA, the
great majority in just five museums, 33% in Colombia and only 12% in European
museums. A total of 1815 species are represented in collections, 95% of those recorded
in the country. Additionally, 9705 and 2855 specimens belong respectively to 180
range–restricted and 102 globally threatened bird species (IUCN categories: CR, EN
and VU). For approximately 80% of collecting localities less than 50 specimens have
been taken, pointing to a historically poor sampling rate in studies of particular
localities. Analysis of density of collections in grids showed that at ≈15 km grain about
80% of the country has had no collecting activity and just 1% of the country has
experienced collection of more than 500 specimens, while at ≈60 km grain only 11% of
the country had collections represented by more than 1000 specimens. Counter–
intuitively, two of the four most intensively collected sites (>3,000 specimens) in
Colombia have subsequently yielded new species for science that had evaded collection
previously. We estimate that possibly more than 100 new bird taxa for science are
pending discovery or description in Colombia. Considering the threats to range–
restricted species, there is an urgent need for continuing ethical and targeted field
inventories, including collecting, across Colombia. It is also important to maintain,
strengthen and co–ordinate the activities of Colombian bird collections. Establishment
of the Darwin–Hernandez database by Project BioMap has created a synthesis of
Verhelst–Montenegro et al.

existing specimen–based information on Colombia’s avifauna, which is available

online to all interested parties to prioritise future research and conservation actions.

Keywords — BioMap; international alliance; biodiversity; birds; collections; Colombia

Colombia is a mega–diverse country (Mittermeier et al., 1997). Its geographical position at the
equator coincides with some of the most topographically complex parts of the Andean Cordillera,
creating a mosaic of different habitats. The country supports an estimated 10 to 20% of the world’s
biodiversity, including approximately 35 000 vascular plants, 733 amphibians, 520 reptiles, 1897
birds and 456 mammals (Chaves & Arango, 1998; Donegan et al., 2012). Colombia is also among the
three countries that hold the greatest number of threatened species and endemic threatened species
(Baillie et al., 2004). Among its birds, 206 species have been listed as threatened or near threatened
by BirdLife International (2013), of which 13 are considered Critically Endangered (CR), 32
Endangered (EN), 69 Vulnerable (VU) and 91 near threatened (NT); according to the International
Union for Conservation of Nature – IUCN criteria (Mace et al., 2008; Birdlife–International, 2013).
In addition, a single bird species has become extinct (EX) in the country, the Andean Grebe –
Podiceps andinus – (Birdlife–International, 2013).
Geographical ranges for many Colombian bird species are still not adequately documented, as
evidenced by the many range extensions and several new species described for science during the last
20 years (Salaman et al., 2001; Salaman et al., 2009a; Donegan et al., 2011; Donegan et al., 2012). To
understand species ranges is important, since they are essential for undertaking threat assessments and
for modelling for inventory and conservation priority setting. Information on geographical ranges can
be obtained from field inventories and surveys, but this is expensive at a large scale, especially in
Colombia where access is difficult to many areas and not possible in others due to political instability.
In contrast, natural history collections and other sources of information such as collectors' notes and
literature are important resources that are relatively easy to access but require synthesising.
For Mexico, Peterson et al. (1998) and Navarro–Sigüenza et al. (2002, 2003) compiled
information from nearly 60 natural history museums and more than 4,000 literature references into a
database containing 450 000 accessions (Peterson et al., 2006). Counts of specimens in 1 × 1 degree
cells, approximately 111 km N–S × 106 km E–W, using a sample of more than 221,000 specimens
from this database, showed that just seven of 240 cells (≈3%) for the country had more than 1,000
collected specimens and could be considered well sampled* (Peterson et al., 1998). Furthermore, the
same authors concluded that collections of the Mexican avifauna are still spatially biased and
incomplete. While to date this extensive dataset has not been made public, its compilers have used it
as a base to conduct a wide range of studies related to the geographical ranges, biogeography and
conservation of the birds of Mexico (Peterson et al., 1999; Peterson et al., 2001; Peterson et al., 2002;
Peterson & Holt, 2003; Peterson et al., 2003; Stockwell & Peterson, 2003; Ortega–Huerta & Peterson,
2004; Peterson et al., 2004; Peterson & Navarro–Sigüenza, 2006; Peterson et al., 2006; Lira–Noriega
et al., 2007; Ramirez–Bastida et al., 2008; Toribio & Peterson, 2008; Peterson & Navarro–Sigüenza,
2009).
Project BioMap, for which data collection was conducted during 2001–2005, constitutes an
international alliance between institutions in the United Kingdom (Natural History Museum – NHM),
Colombia (Instituto de Ciencias Naturales, Universidad Nacional de Colombia – ICN) and the USA
(Conservation International – the Center for Applied Biodiversity Science – CABS), funded largely
by the UK government’s Darwin Initiative and by Conservation International. It aimed to improve
avian biodiversity knowledge in Colombia through data sharing of information relating to all
Colombian bird specimens accessible in the natural history museums around the world. This paper
reviews the development of the project and, using the data assembled by it, presents an overview of
the history and completeness of bird collections from Colombia .

*
The authors set this threshold assuming that in a sample of 1000 specimens may be represented at least 100
species, which is very likely the minimum number of species found in any local avifauna in Mexico.

Notulae Ornitologicae Columbianae 2 3

 Project BioMap – Colombia

Methodology

Data capture, compilation and checking. — Project BioMap began in 2001 and first made captured
data publicly available on the Internet in 2006, a process that can be conveniently divided into five
stages (Fig. 1). Prior to museum visits being undertaken, curators were contacted in order both to
obtain their agreement to involvement and to discover their estimates of the numbers of bird
specimens from Colombia in each collection, thereby enabling production of a realistic work
timetable and prioritisation of data capture in museums with larger numbers of Colombian specimens.
From January 2002 until December 2003, BioMap team members, generally in pairs, visited targeted
collections in each major region holding significant numbers of Colombian specimens (i.e. Colombia,
North America and Europe). When a particular museum had electronic data on its collections, this
was obtained in advance and was prepared and organised in either Access or Excel formats to
leverage past efforts. Where pre–existing electronic data were unavailable, specimen data were
directly entered from the museum catalogues using a BioMap entry tool. A standard data entry
protocol was used by the project to ensure as far as possible coherence in data structure.

Preparation
Data entry tool developed, curators and collection managers of targeted museums
contacted, museum holdings estimated and visits prioritised

Data capture
Museums' catalogue and available electronic datasets
captured and, where possible, verified against specimens

Datasets compilation
Verified museum datasets standardised
and migrated to BioMap Access relational database

Database revision
Database master tables and fields homogenised and synchronised in agreement with the data
entry protocol

Publication on the Internet

Data uploaded to server, Google Earth localities interface developed, login system for clients
administration established and feedback by end users integrated
Figure 1. Main stages of the Project BioMap effort to catalogue and verify Colombian bird specimens.

The data entry tool was developed by CABS staff and written in Delphi language, and the local
database implemented in Microsoft Access 2000. The tool consisted of an interface with an initial
window where the name of the cataloguer was entered, followed by a second window where data
input was performed. All the information digitised and verified during collection visits was compiled
Verhelst–Montenegro et al.

and curated into a unique relational database, the ‘Darwin–Hernandez* database’. The database
contains a total of 105 fields and comprises one main table ‘RegMus_Especimen’, which contains all
the accessions and retrieves information from 11 satellite tables (Fig. 2).
The information fields included in the database were classed as obligatory, desirable or
optional. Fields were designed to register information related to the museum (e.g. museum name,
acronym), existing museum database references (catalogue number), the taxonomy of the specimen
(i.e. family, genus, species and subspecies) and the collection event (e.g. locality, date, collector, field
number, etc.), as well as age, sex, general notes and information about the existence of accessory data
on the label (i.e. soft parts, DNA, gonads state, body measurements, weight and fat) together with, if
required, other comments.
Five dictionaries were used in the data entry window: 1) taxonomy, 2) localities, 3)
organisations, 4) contacts and 5) bibliography. The taxonomy dictionary was initially based on
Salaman et al. (2001), but including amendments suggested by F.G. Stiles, curator of birds at ICN.
This dictionary included a complete guide of synonyms. Subsequently, after the compilation of the
database, it was revised and updated to follow most of the genus and species level changes adopted by
Remsen et al. (2007). The localities dictionary was based on the ornithological gazetteers of
Colombia by Paynter (1997) and Paynter & Traylor (1981), but updated and kept under review using
online gazetteers and such other sources as printed maps (scales 1:100 000 to 1:500 000) from the
‘Instituto Geografico Agustin Codazzi’ (IGAC). The organizations and contacts dictionaries had
respectively lists of all the institutions and collections with Colombian specimens and the contact
details of the curator or the collection manager. Finally, the bibliography dictionary had a list of
relevant published literature. However, this dictionary was not used in data capture, since data from
the literature were not included in the database.

Figure 2. Table relationships in the Darwin–Hernandez database.

*
Honouring both Charles Darwin and the Colombian naturalist Jorge Ignacio Hernandez–Camacho, who led
collecting activity and environmental research in the country during five decades until his sudden death in 2001.

Notulae Ornitologicae Columbianae 2 5

 Project BioMap – Colombia

Once the dataset was prepared or data from available museum catalogues had been entered
using the data entry tool, cataloguers checked each record against the actual specimen. Revisions were
made to ensure the dataset correctly listed the identity of each specimen (to subspecies where
possible) and other information written on its attached labels, especially locality. Furthermore, it was
possible to check that each specimen still existed in each collection, and whether it was loaned or
damaged. The information was then compiled into a single database in Microsoft Access version
2003.
After all datasets were compiled, locality and taxonomic dictionaries were verified and a
directory of collectors was created. This revision was designed to homogenise names in the
department, municipality and locality fields in the locality table and names in family, genus, species
and subspecies fields in the taxonomy table and check data were in the correct fields after migration
of each dataset to Access. Georeferencing was undertaken through consultation of various gazetteers
and other sources, including Conservation International’s unpublished database of localities for
amphibians and mammals, the Red Book of the Birds of Colombia (Renjifo et al., 2002), Alexandria
Digital Library Gazetteer (ADL, 2004) and the National Geospatial Intelligence Agency Geographical
Names Server Database (NGA-GNS, 2004). The original datasets as they existed before compilation
were kept as backups of the information by Conservation International – Colombia.
Once the compilation and revisions were completed, the database was uploaded to the BioMap
internet portal (http://biomap.net) in 2006 for public use. After a simple process of registration,
including accepting terms and conditions of data use, and approval, any user may access the data for
consultation.

Assessment of spatial distribution of collected specimens. — In order to assess the spatial

distribution of collected specimens, as part of various preliminary analyses conducted by Verhelst
(2011), all accessions in the Darwin–Hernandez database were once more revised, corrected and
filtered; this resulted in an updated version of the database, not available yet on–line, used in analyses
presented here. First, using DIVA–GIS (Hijmans et al., 2005), we revised and, whenever necessary,
corrected georeferencing using different gazetteer resources (Paynter & Traylor, 1981; Paynter, 1997;
ADL, 2007; NGA–GNS, 2007). Next, all records with uncertain localities (i.e. no georeferencing,
localities approximated to ‘department’ and ‘Bogota’ trade skins), with incomplete taxonomy to
species level and with non–continental localities (e.g. islands of Malpelo, San Andres and
Providencia) were deleted. Afterwards, the data were imported into Arc View 3.3 and, for each
specimen, we extracted data on the elevation of each collecting locality using a Digital Elevation
Model* at ≈1 km (Jarvis et al., 2004) and the Grid Analyst extension.
The new dataset was exported to Microsoft Excel and, for each accession, two new columns
were added showing the upper and lower altitudinal limits for the species, based upon Stotz et al.
(1996) and Hilty & Brown (2001). To retain only records of species agreeing with the altitudinal
range reported in the literature or possible expansions of the altitudinal known range, records were
kept when the extracted altitude was within an arbitrary limit of 500 m above and below the range of
altitudes noted in the literature. Records outside the specified range, which were potentially errors,
were filtered out from the database. Following these filters, we retained 174 330 specimens that had
complete locality and taxonomy data. The final dataset was exported as a text file and imported into
DIVA–GIS as shape file. Once in DIVA–GIS, the simple point–to–grid conversion procedure was
used to estimate the number of collected specimens in grids at ≈15 km and ≈60 km over the country.

Results and analyses

Location of Colombian bird specimens. — We were permitted to obtain information from 89

museums in 19 countries that house a total of 218 194 Colombian bird specimens (see Appendix S1 in
online Supplementary Materials, which includes a list of the museums and their holdings). The

*
Aggregated (expanding grid and ignoring no data) from the void–filled seamless SRTM DEM data
V1 at ≈90 m resolution.
Verhelst–Montenegro et al.

Darwin–Hernandez database holds information on all of these, with exception of 143 specimens
deposited in the Cleveland Museum of Natural History and 392 specimens deposited in the Museum
and Institute of Zoology–Polish Academy of Sciences, which due to technical issues were not
included in this version of the database. Consequently, the current version of the database contains
only information on 217 659 specimens. Otherwise, Project BioMap includes every collection except
one worldwide that we are aware contains more than a negligible number of Colombian specimens.
The exception is Instituto Alexander von Humboldt – IAvH (previously known as the INDERENA
collection), in Colombia, which reportedly holds 13 411 further specimens (IAvH, 2011), giving a
total of 231 605 bird specimens worldwide that have been collected in Colombia.
The number of collections containing Colombian bird specimens (IAvH included) was similar
between the western and eastern hemispheres, with 43 and 46 museums respectively. However,
collections in Colombia and the USA accounted for 88% of all Colombian specimens (Fig. 3a).
Furthermore, 71% of all specimens are held in just nine collections – five in the USA, three in
Colombia and one in the UK (Fig. 3b) – that together comprise the core resource for taxonomic
research concerning Colombian birds.

a. b.
ICN
Other countries 14%
United Kingdom
3% 5%
Germany
4% NMNH
12%
Other museums
29%

Colombia United States IAvH

33% 55% 6%
AMNH
NHM 11%
3%

MLS
4% FMNH ANSP
CMNH 9%
6% 6%
Figure 3. Percentage of bird specimens from Colombia deposited in (a.) different countries, and (b.) different
museums collections. Major museums in Colombia: Instituto de Ciencias Naturales (ICN); Universidad de La
Salle (MLS); and Instituto de Investigacion de Recursos Biologicos Alexander von Humboldt (IAvH). Major
Museums in the USA: National Museum of Natural History (NMNH); American Museum of Natural History
(AMNH); Academy of Natural Sciences of Philadelphia (ANSP); Carnegie Museum of Natural History
(CMNH); and Field Museum of Natural History (FMNH). Major museum in United Kingdom: The Natural
History Museum (NHM).

Species represented. — The Darwin–Hernandez database contains information on 1815 species that
belong to 89 families and 663 genera. The best–represented families are: tyrant flycatchers
(Tyrannidae, 190 species), hummingbirds (Trochilidae, 187 species), tanagers (Thraupidae, 151
species), ovenbirds (Furnariidae, 105 species), antbirds (Thamnophilidae, 104 species), sparrows
(Emberizidae, 57 species), parrots (Psittacidae, 55 species), wood warblers (Parulidae, 53 species) and
hawks and eagles (Accipitridae, 51 species). Those nine families account for just over 50% of the bird
species contained in the database (Fig. 4). In this context, it should be noted that the database’s
taxonomic table has not been updated since 2007 to take account of the most recent changes adopted
by the South American classification Committee of the American Ornithologists’ Union (Remsen et
al., 2013).

Notulae Ornitologicae Columbianae 2 7

 Project BioMap – Colombia

Tyrannidae
10%
CR
9%
Trochilidae
10%

Other families
48% Thraupidae
8% EN
29%
Furnariidae
6%

Thamnophilidae
6%
Emberizidae VU
3% 61%
Accipitridae
3% Psittacidae EX
Parulidae
3% 1%
3%

Figure 4. Proportion of species by family Figure 5. Species categorised as threatened in

represented in the Darwin–Hernandez the Darwin–Hernandez database, distributed
database. by category.

The Darwin–Hernandez database contains data for 9705 specimens of 180 range–restricted
species (97% from the total) found in Colombia (Stattersfield et al., 1998; Stiles, 1998), as well as
data for 2855 specimens of 102 species (89% from the total) categorised as threatened (Birdlife–
International, 2013) (Fig. 5).
The database has further revealed a number of previously overlooked new species records for
the country, including Leach's Storm–Petrel (Oceanodroma leucorrhoa), Blue–mantled Thornbill
(Chalcostigma stanleyi), Brown–banded Puffbird (Notharchus ordii), Yellow–throated Warbler
(Dendroica dominica) and Black–and–white Tanager (Conothraupis speculigera) (Donegan et al.,
2009; Salaman et al., 2009a; Lobo–y–HenriquesJC et al., 2012).

Collection localities. — When the database was first compiled in 2004, a total of 8135 collection
localities were generated. This initial list included many duplicates and errors in the coordinates, and
approximately 50% of the localities did not have any coordinates assigned. After almost a year of
revisionary work during 2004–2005, the list was reduced to 3866 collecting localities. Of these, only
167 were not assigned coordinates, either because it was not possible to assign any, such as for
example the ‘Bogota’ skins, for which the recorded locality represents the place from which they were
shipped rather than collected (Rasmussen & Prys–Jones, 2003), or because it was not possible to find
the locality in gazetteers or the literature. Possibly up to a further 570 localities may still be
duplicates, i.e. localities with different names that were assigned the same pair of coordinates. These
resulted from approximation of specific localities not georeferenced in any gazetteer or map to the
most certain nearest locality. Although the most labour–intensive effort by the project was the
revision of each specimen record, we underestimated the effort required for locality data checking and
georeferencing, which continued to be refined later within other studies that made use of the database
(Verhelst, 2011). Unfortunately, due to lack of funding this new version has not yet been uploaded to
the internet and a further revision of the data has not been possible.
Results show that only small numbers of specimens have been collected at most localities. For
about 80% of localities, less than 50 specimens have been collected (Fig. 6). Given both that each
specimen does not necessarily correspond to a different species and that in almost all localities of
Colombia, including even those with severely fragmented habitats such as the coffee region in the
Colombian Andes, more than 100 species can be recorded within a few days (Botero et al., 1999), this
Verhelst–Montenegro et al.

demonstrates that for most localities available museum collections are well below the minimum
necessary to detail the avifauna present. Furthermore, it is important to note that a locality of
collection, represented by a unique pair of coordinates, does not necessarily represent the exact place
where the specimens recorded for it were collected, but only an approximation. Many collectors
would have had base camps around which they performed their activities, and it is probable that some
georeferenced localities in fact include specimens taken up to a few kilometres away.

3,500
3,168

3,000
Number of localities (n)

2,500

2,000

1,500

1,000

500
250
196 163
40 18 7 13 9 2
0
< 50 < 100 < 200 < 400 < 600 < 800 < 1,000 < 2,000 < 4,000 > 4,000

Specimens per site locality intervals (–)

Figure 6. Frequency distribution of localities in relation to number of collected specimens.

On a coarser spatial scale, maps of the density of collections at ≈15 and ≈60 km grain further
demonstrate that a major proportion of the country has either no or very few and sparse collections
taken within it. At ≈15 km grain, more than 80% of the cells in the country have had no collecting at
all, while only 1% of Colombia has experienced collection of more than 500 specimens (Figs. 7a,b).
At ≈60 km grain, only 11% of the cells have had collections totalling more than 1000 specimens
(Figs. 7c,d). Further analyses conducted by Verhelst (2011), counting species richness and effort in
grids using DIVA–GIS, demonstrated that at both grains there is a strong relationship between
richness and collection effort (Fig. 8), with no evidence of an asymptote of species being reached
even for localities at which large numbers of specimens have been collected.
Overall, the results confirm that most collections have been made in the western half of the
country, mainly in the Andes, some parts of the Pacific coast, and around ‘Sierra Nevada de Santa
Marta’ in the north. In particular, much collecting activity has been focused in the vicinity of major
cities, such as ‘Bogota’, ‘Medellin’, ‘Cali’, ‘Ibague’, ‘Manizales’, ‘Popayan’, ‘Bucaramanga’,
‘Cucuta’, ‘Pasto’, ‘Armenia’, ‘Pereira’, ‘Monteria’, ‘Santa Marta’, ‘Riohacha’, ‘Quibdo’,
‘Villavicencio’, ‘Florencia’ and ‘Mocoa’.
By contrast, there has been a remarkably poor collecting effort in the Orinoco and Amazon
regions. Sixty years ago, Meyer De Schauensee (1948-1952) pointed out how unexplored those areas
were at that time, asserting ‘…when comprehensive collections are made on the Amazon, the Vaupes,
the Rio Negro, and along the Orinoco and Meta Rivers, a conservative estimate would add 200 forms
to the Colombian Avifauna…’. Although since then more than 600 additional species have been
documented and added to the list of birds of Colombia, some resulting from splits but many being
new records, the results presented here demonstrate that the birds of eastern Colombia remain poorly
known.
There are also still many areas west of the Andes, in the Caribbean, Pacific and even Andean
regions, where collections have been absent or small, and there is a clear and present need for further
studies (Fig. 7). Evidence of the generally poor state of knowledge regarding Colombian birds is
demonstrated by the rate of recent discoveries despite the intense political instability in Colombia. For

Notulae Ornitologicae Columbianae 2 9

 Project BioMap – Colombia

a. c.

b. d.

Figure 7. Density of collections across Colombia at ≈15 km (a., b.) and ≈60 km (c., d.) grain.
Verhelst–Montenegro et al.

300

Richness (No. of species)

250

200

150

100

50

0
0 2,500 5,000 7,500 10,000 12,500 15,000 17,500 20,000
Effort (No. of specimens)

S15 S30 S60 S120

Figure 8. Richness versus effort (No. of collected specimens) at different grain (≈15–120 km), modelled using
the species–area relationship (logarithmic model). In all cases both variables showed a very strong relationship
(R2 > 0.95 p < 0.05). Graph reproduced and slightly modified from Verhelst (2011).

example, since the landmark publication of Hilty & Brown (1986), 171 bird species new to Colombia
had been reported by 2009 (Salaman et al., 2009a) and another 27 new species records were
documented by 2012, totalling 1897 bird species (Donegan et al., 2012).
Importantly, the rate of new bird taxa for science described during the last 20 years as result of
new explorations in the field has accelerated (Fig. 9), with 20 new species and subspecies described
from 1986-2009 (Salaman et al., 2009a) despite often complicated fieldwork conditions. New species
for science from Colombia include: Cloud–forest Pygmy–owl Glaucidium nubicola (Robbins &
Stiles, 1999), Chiribiquete Emerald Clorostilbon olivaresi (Stiles, 1996), Bogota Sunangel
Heliangelus zusii (Graves, 1993), Gorgeted Puffleg Eriocnemis isabellae (Cortes–Diago et al., 2007),
Parker’s Antbird Cercomacra parkeri (Graves, 1997), Cundinamarca Antpitta Grallaria kaestnaeri
(Stiles, 1992), Urrao or Fenwick’s Antpitta Grallaria fenwickorum (Barrera et al., 2010; Caranton &
Certuche, 2010), Upper Magdalena Tapaculo Scytalopus rodriguezi (Krabbe et al., 2005),
Stiles’ Tapaculo Scytalopus stilesi (Cuervo et al., 2005), Chestnut–capped Piha Lipaugus weberi
(Cuervo et al., 2001), Antioquia Bristle–tyrant Phylloscartes lanyoni (Graves, 1988), Antioquia Wren
Thryophilus sernai (Lara et al., 2012), Munchique Wood–wren Henicorhina negreti (Salaman et al.,
2003), Choco Vireo Vireo masteri (Salaman & Stiles, 1996) and Antioquia Brush–finch Atlapetes
blancae (Donegan, 2007).
The most intensively sampled ‘locality’ was ‘Bogota’ with 15 877 specimens, including
specimens labelled ‘Vicinity of Bogota’, ‘Sabana de Bogota’, ‘Bogota Uncertain Locality’, etc.
(Fig. 10). The next ‘localities’ with most specimens are ‘Colombia’ (not including New Granada) and
‘No Locality Data’ (Fig. 10). The best represented actual localities, with >2000 specimens each, are:
‘Bonda’ (‘Magdalena’), ‘Guayacana’ (‘Nariño’), ‘Munchique’ (‘Cauca’), ‘Villavicencio’ (‘Meta’),
‘Popayan’ (‘Cauca’), ‘Morelia’ (‘Caqueta’), and ‘Ricaurte’ (‘Nariño’).
In addition, from a total of 16 579 specimens that have uncertain localities (e.g. Bogota
Uncertain Locality, Colombia Uncertain Locality, New Granada) or Bogota as locality, we estimate
that some 10 179 are likely to be ‘Bogota trade skins’ since the accessions have notes regarding the
locality that point in that direction. These include 2929 specimens clearly stating ‘Bogota trade skin’,
‘native skin’, ‘native collection’, ‘native Bogota Collection’, etc., 1348 specimens collected prior to
1900, 5335 specimens having no date or collector, and 571 specimens that have clearly passed

Notulae Ornitologicae Columbianae 2 11

 Project BioMap – Colombia

Figure 9. Type localities for some new bird taxa described for science during the last 20 years in Colombia in
relation to areas in which collections are equal or > 1,000 specimens at ≈60 km grid scale.

SAN AGUSTIN

FUSAGASUGA=FUGAZUGAZA
AGUADA = LA AGUADITA = AQUADITA
JURADO, RIO
SANTA MARTA

MEDELLIN
LAGUNETA
CANDELA = LA CANDELA

SAN ANTONIO
Localities

SAN ANTONIO
CALI
EL TAMBO

RICAURTE

LA MORELIA = MORELIA = LA MURELIA

VILLAVICENCIO
POPAYAN

MUNCHIQUE, CERRO (1)

GUAYACANA = LA GUAYACANA
[BOGOTA, UNCERTAIN LOCALITY]
BONDA
[NO LOCALITY DATA]

[COLOMBIA, UNCERTAIN LOCALITY]

BOGOTA

0 2 000 4 000 6 000 8 000 10 000 12 000 14 000

Specimens collected

Figure 10. Site localities most intensively collected in Colombia. Note that localities in brackets are artificial
constructs and not equivalent to the others. Moreover, it may be that some specimens with locality Bogota may
correspond to Bogota trade skins, but this needs further assessment.

Notulae Ornitologicae Columbianae 2 12

Verhelst–Montenegro et al.

through the hands of dealers, e.g. ‘Verreaux collection’, ‘Lafresnaye collection’, ‘Maximillian
collection’, etc.

Main collectors and collections over time. — Approximately 23% of specimens (50 426) do not have
any data concerning the collector. Most of these were Bogota or Colombia trade skins and other old
specimens. The number of early collectors is almost certainly underestimated, because specimens
taken prior to the 1900s are less likely to have labels showing collector name.
By far the most important known collectors in Colombia were M. A. Carriker Jr. and K. Von
Sneidern, who each collected about 28 000 specimens (Fig. 11). Other major bird collectors for the
country, who each took more than 3 000 specimens, were: P. Bernal, Brother Niceforo Maria, J. I.
Borrero, W. B. Richardson, A. A. Allen, F. C. Lehmann and Father A. Olivares (Fig. 11).
The temporal distribution of specimens collected provides insight into the history of
ornithological collecting efforts in Colombia. In total the Darwin–Hernandez database holds 165 420
(76%) specimens with information regarding their year of collection. Additionally, 13 947 more
specimens deposited in the CM, which have a date in the original files, do not hold entries in the
Darwin–Hernandez database because of a failure to import the corresponding fields from the original
file due to unknown technical reasons to us. However, these were used too to calculate the total of
collected specimens reported in the following analyses. The first extant bird specimen with known
date is an individual of the Oleagineous Hemispingus (Hemispingus frontalis, AMNH catalogue
number 511466, Colombia) collected in 1773, originally from the R. De Dalmas collection. From the
early 1800s, the number of specimens collected per 25–year period increased consistently up to a
maximum of over 60 000 in the period from 1950–1974, before more than halving during the
subsequent 25 years (Fig. 12a). Contrastingly, collecting trends change more unpredictably when
assessed using a finer 10–year window (Fig. 12b). This reveals a first, minor peak in the last decade of
the 19th century. Otherwise, during the past century, two major collecting peaks coincided with each
World War and immediate post–war periods, although the latter peak continued through the 1960s
(Fig. 12b). It is important to highlight that inclusion of IAvH collection data might change the shape
of the curve of specimens held in Colombian museums, notably in inflating values slightly in the
period 1940–2004.
In the last quarter of the 20th century, overall collecting activity decreased primarily because
foreign institutions reduced and later stopped collecting activities in Colombia and ornithologists
increasingly used other techniques, such as field observations, mist–nets, photography and sound
recordings, to document records. Additionally, in 1959 the National Institute of Renewable Natural
Resources and Environment (INDERENA) was created, as part of the Ministry of Agriculture, to be
in charge of the research, management and protection of the country’s natural heritage. This first
effort was strengthened with the publication of the ‘Law 2811’ or ‘National Code of Renewable
Natural Resources and Protection of the Environment’ in 1974, which amalgamated isolated
legislation related to the environment in the country since the beginning of the 20th century, becoming
the keystone of environmental policy in Colombia (Sanchez–Perez, 2002). This certainly also
contributed to the decreased export of bird skins from the country.
Prior to 1920, a total of 47 037 specimens were collected. The 1890s were an important period
(8591 specimens), with key collectors being G. H. Hull (2087 specimens), W. W. Brown Jr. (1903
specimens), H. H. Smith (1054 specimens) and J. H. Batty (520 specimens). This material is now
mainly at AMNH (4306 specimens) and the MCZ (2069 specimens). During the 1910s, significant
collections (31 415 specimens) were compiled by M. A. Carriker Jr. (15 185 specimens), W. B.
Richardson (3512 specimens), A. A. Allen (3434 specimens), L. E. Miller (2276 specimens), F. M.
Chapman (1291 specimens), M. Gonzalez (820 specimens), G. K. Cherrie (814 specimens), Brother
Niceforo Maria (565 specimens) and H. Boyle (473 specimens), the great majority of which went to
the AMNH (13 029 specimens), CM (12 735 specimens) and the ANSP (1901 specimens). Almost
half of these specimens were collected on AMNH expeditions.
In the 1940s, collecting activity peaked again with 33 749 specimens taken by foreign and
national collectors. Among them K. Von Sneidern and M. A. Carriker Jr. collected 5017 bird
specimens for the ANSP, whilst A. Wetmore collected 825 specimens for the NMNH, both museums
in the USA. A major part of the increased activity during this decade was the result of the creation of
several natural history museums in Colombia, which began to assemble substantial collections that

Notulae Ornitologicae Columbianae 2 13

 Project BioMap – Colombia

Wheeler, T.H.
Sciama
Chapman, F.M.
Hno. Serna, M.A.
Marinkelle, C.J.
Salmon, T.K.
Haffer, J.
Romero, H.
Mena, T.
Brown, W.W., Jr.
Collectors

Smith, H.H.
Miller, A.H.
Palmer, M.G.
Hull, G.H.
Miller, L.E.
Olivares, A.
Lehmann, F.C.
Allen, A.A.
Richardson, W.B.
Borrero, J.I.
Hno. Nicéforo, M.
Bernal, P.
Von Sneidern, K.
Carriker, M.A., Jr.

0 3 000 6 000 9 000 12 000 15 000 18 000 21 000 24 000 27 000 30 000

Specimens collected

Figure 11. Most important bird collectors in Colombia.

a.
70 000
65 000
60 000
55 000
50 000
Specimens collected

45 000
40 000
35 000
30 000
25 000
20 000
15 000
10 000
5 000
0
≤ 1849 ≤ 1874 ≤ 1899 ≤ 1924 ≤ 1949 ≤ 1974 ≤ 2004

Years intervals

b.
33 000
31 500
30 000
28 500
27 000
25 500
24 000
22 500
Specimens collected

21 000
19 500
18 000
16 500
15 000
13 500
12 000
10 500
9 000
7 500
6 000
4 500
3 000
1 500
0
≤ 1849

≤ 1859

≤ 1869

≤ 1879

≤ 1889

≤ 1899

≤ 1909

≤ 1919

≤ 1929

≤ 1939

≤ 1949

≤ 1959

≤ 1969

≤ 1979

≤ 1989

≤ 2004

Years intervals
Specimens Colombia Specimens United States Specimens Europe

Figure 12. Temporal distribution of bird collection in Colombia using two different window sizes: a) 25–year
window, and b) 10–year window; with collecting activity sub–divided according to whether specimens ended up
in Colombian, USA or European museums. Please note that the last interval in each figure is irregular adding
respectively five more years in each case, thus using all information available in the database.

Notulae Ornitologicae Columbianae 2 14

Verhelst–Montenegro et al.

remained within the country. However, it is important to highlight that the number of specimens
collected by M. A. Carriker Jr. and A. Wetmore during this and the next decade for the NMNH may
well be underestimated in the current version of the database, because of a failure to import the
collectors field from the original files due to technical reasons unknown to us.
In 1939, the Instituto de Ciencias Naturales (ICN) was established as part of Universidad
Nacional in Bogota and quickly became the ‘cradle’ of Colombian ornithology. Some of the
ornithologists trained in ICN moved to Cali and Popayan in southwest Colombia, where they started
and strengthened local collections and ornithological groups attached to public universities (Naranjo,
2000). Some remarkable Colombian ornithologists and naturalists who started their activities during
this decade and continued later included Brother Niceforo Maria (1274 specimens), who although
born in France lived for 71 years in Colombia (Tamsitt, 1980), F. C. Lehmann (974 specimens), J. I.
Borrero (809 specimens), C. Herrera (284 specimens), A. Dugand (129 specimens) and Father A.
Olivares (98 specimens). Their collections during this decade went mainly to the ICN, IAVH
(information not included), Universidad de La Salle and Universidad del Cauca.
During the 1950s, the most important collectors were M. A. Carriker Jr. (6093 specimens) and
K. Von Sneidern (4655 specimens), whose material ended up mainly in the FMNH (5301 specimens)
and the NHMLAC (3453 specimens). The principal Colombian collectors active in this decade were
Brother Niceforo Maria (2500 specimens), Father A. Olivares (2158 specimens), F. C. Lehmann
(1475 specimens), J. I. Borrero (1299 specimens), P. Bernal (521 specimens) and J. I. Hernandez–
Camacho (199 specimens).
The 1960s marked an important change in the history of ornithology for Colombia, as
specimens secured by local institutions began to represent the major proportion of the total collected.
The most important collectors for this decade were P. Bernal (4559 specimens), M. A. Carriker Jr.
(3168 specimens), K. Von Sneidern (2049 specimens), J. I. Borrero (1536 specimens), J. Haffer (998
specimens), C. J. Marinkelle (947 specimens), A. Pazos (865 specimens), Father A. Olivares (595
specimens), P. Orejuela (478 specimens) and G. Cataño (365 specimens). ICN was the museum that
received most of the Colombian material taken during the decade (8842 specimens).
In the 1970s, the principal collectors included K. Von Sneidern (3232 specimens), P. Bernal
(1816 specimens), J. I. Borrero (821 specimens), N. Moreno (839 specimens), H. Romero (673
specimens), G. Cataño (584 specimens) and Brother M. A. Serna (535 specimens). In the last two
decades of the 20th century the most important collectors were H. Romero (667 specimens), G.
Arango (506 specimens), and A. Negret (507 specimens). Other important collectors under–
represented in the Darwin–Hernandez database, but held by IAVH, include J. I. Hernandez–Camacho,
J. V. Rodriguez–Mahecha (on collecting expeditions with INDERENA, mostly in the 1970s–1980s)
and M. Alvarez–Rebolledo, A. M. Umaña and M. Sierra (on collecting expeditions of IAvH in the
1990s to 2009).

Concluding remarks

Project BioMap represents a model of international ornithological cooperation and data sharing and
dissemination that is already greatly assisting our understanding of the avifauna of Colombia and that
demonstrates the importance of existing biological collections. The database contains the data from
94% of the Colombian bird specimens we are aware of, which has been entirely verified,
georeferenced as far as possible and is available online. We hope that this tool will aid researchers
working with Colombian birds long into the future. To December 2011, 526 registered users have
accessed the Darwin–Hernandez database (http://biomap.net), using its search engine and locations
linked to Google maps. The data have been cited and used in a number of recent studies involving
descriptions of new taxa (Donegan & Huertas, 2006; Donegan, 2007, 2008; Donegan & Avendaño–C,
2008; Salaman et al., 2009b) and in biogeographical research (Graham et al., 2010; Verhelst, 2011;
Velasquez-Tibata et al., 2013). It is important to note that the Darwin–Hernandez database does not
replace the need to visit collections and inspect specimens or photographs of them. However, it
provides a tool to reveal the extent of material available for consultation and encourages researchers

Notulae Ornitologicae Columbianae 2 15

 Project BioMap – Colombia

to plan their research itineraries and visit important collections to examine material first–hand rather
than to visit museums on a speculative basis.
Despite continued collecting efforts conducted by national institutions in Colombia such as the
ICN and IAvH, among others, there has been a sharp decrease in collecting during the past three
decades in comparison to earlier decades of the 20th century. This not only corresponds to a period of
political instability in rural areas of the country, but also to the increased usage of other ornithological
fieldwork methods (e.g. banding or ringing, sound–recording, photography and observations).
Furthermore, use of collecting of living specimens as a fieldwork technique has become increasingly
controversial. Some leading ornithologists advocate a continuation of past mass collecting efforts to
extend and consolidate national reference collections (e.g. Cuervo et al., 2006). On the other hand,
others have raised concerns about the ethics of collecting large numbers of bird specimens. In their
view, collecting of bird skins is no longer necessary for the documentation of records or many other
aspects of ornithological study (e.g. Donegan, 2008). There are many trained and active bird
collectors in Colombia, but most focus collecting on specific individuals that represent new or
interesting material rather than aiming for quantity. Records of birds also continue to be documented
and georeferenced in new ways. For example, ProAves Foundation (www.proaves.org) has databases
containing nearly 300 000 accessions detailing observations and mist–net captures with biometric and
complete locality data. Similarly, the Birdwatchers National Network of Colombia (RNOA–
Colombia) has a georeferenced database with nearly 400 000 records (http://rnoa.org/) on bird
observations along the country. In addition, several Colombian ornithologists have uploaded
thousands of georeferenced sound recordings in www.xeno-canto.org or have deposited them at
IAVH's sound archive, and various online sites for the sharing of photographic records exist (e.g.
flickr). When combined with specimen data from Project Biomap, these resources together form a
formidable data set that can be used to assess species distributions, ecology and taxonomy.
The spatial paucity of bird collections across Colombia is noteworthy, especially given the
tremendous loss of natural habitat across the Andes, where the majority of range–restricted species are
concentrated. There remain many gaps in our knowledge, illustrated in recent years by two new bird
species being discovered at Munchique (Salaman et al., 2003) and near Bonda, Santa Marta, within
1 km of Carriker’s old home (N. Krabbe in litt.) – two of the four most intensively collected sites in
Colombia (>3000 specimens each). However, both new species had evaded detection before the 1980s
and only recently was enough information collected to confirm their status as new taxa.
Considering the increasing rate of new bird taxon descriptions and the very uneven collecting
effort across the country, we conservatively estimate that possibly 100 new bird taxa for science await
discovery or description in Colombia during the next 100 years. Considering the threats to range–
restricted species, there is an urgent need to undertake targeted fieldwork, including targeted ethical
collecting, across Colombia, particularly in parts of the Eastern, Central and Western Cordilleras and
their interior montane valleys, as well as the Choco region.
It is also very important to maintain and strengthen existing collections and ornithological
fieldwork capacity within Colombia and increase cooperation and coordination between different
Colombian organisations. Project BioMap has established a unique opportunity to better understand
the avifauna of Colombia, to prioritise research and to assist conservation actions in the future.
However, many challenges still lie ahead to continue improving the database georeferencing and
correcting some already known errors that exist in the database. At the moment we estimate that about
1 to 10% of the accessions imported from the different museum datasets became corrupted at some
point in the process. Particularly, we have detected problems with data from the Western Foundation
of Vertebrate Zoology (WFVZ), which needs further revision and to be imported again in the
database. These challenges await new proposals and funding, which have yet to be secured.

Acknowledgements

Project BioMap was made possible thanks to the following staff: A. Morales, D. Arzuza,
C. I. Bohorquez, N. Cleere, S. De la Zerda and L. Rosselli, together with the supervision and
suggestions of G. F. Stiles. We would like to thank very strongly the curators and staff of all the

Notulae Ornitologicae Columbianae 2 16

Verhelst–Montenegro et al.

natural history museums visited during the development of the project, as well as the staff at the
Natural History Museum (J. Stewart) and Conservation International – Colombia (J. V. Rodriguez, A.
Espinel, F. Arjona, and D. Estepa) who contributed greatly towards it. This project was funded by the
UK Darwin Initiative – DEFRA, Conservation International – CABS and CI – Colombia, as well as
by two fellowships from each of the AMNH, the ANSP and King’s College London. We are indebted
to M. Mulligan, who supervised both Darwin Fellows (C. I. Bohorquez and J. C. Verhelst) while they
were at King’s College London completing their Masters degrees as part of Project BioMap, and T.
Donegan, who provided useful comments on this paper. We would like to thank especially the many
people in the USA, Europe and Colombia, including museum staff and friends, who collaborated with
accommodation, transportation, and moral support during data capture, since without their support
this project would not have succeeded. Finally, we would like to thank Z.–Q. Zhang, T. Chesser who
made valuable comments on the final versions of this manuscript .

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