Saboya Acosta Urbina Cardona 2023 Current State of Knowledge of Paramo Amphibians in Colombia Spatio Temporal Trends

Review Article
Tropical Conservation Science

Volume 16: 1–23
Current State of Knowledge of Páramo © The Author(s) 2023
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Amphibians in Colombia: Spatio Temporal sagepub.com/journals-permissions
DOI: 10.1177/19400829231169984
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Trends and Information Gaps to Be
Strengthened for Effective Conservation
Liliana Patricia Saboyá Acosta1  and J. Nicolás Urbina-Cardona2
Abstract
Background and Research Aims: Globally, Colombia is the country with the largest extent of Páramos (delimited in 36
complexes) and with the greatest number of amphibian species in this ecosystem. This work consolidated scientific literature on
the amphibians of the Colombian Páramos to characterize temporal, taxonomic, thematic, and geographic patterns, which allow
us to identify information gaps that must be fulfilled to achieve effective species conservation. Methods: We conducted a
systematic literature survey with seven different search strategies and generated a database. We read each document’s
Abstract, Methods, Study Area, Results, and supplementary material, following the PRISMA (Preferred Reporting Items for
Systematic reviews and Meta-Analyses) protocol. Results: We found 405 documents published between 1863 and 2021. The
composition and richness of 142 amphibian species (95 endemics to Colombia), presented significant differences in Páramo
complexes and between sectors. Since 2000, the diversity of research topics has increased with a high proportion of studies on
Natural History, Systematics and Taxonomy, and Conservation, distributed between 19 and 22 of the departments with
Páramos in their jurisdiction. However, much of this knowledge concentrates in less than 20% of total species in just 6% of
Páramos complexes. Conclusion: We found critical shortfalls in taxonomy, spatial information, and conservation actions on
Páramos amphibians. We need to increase studies that include field data in more geographic areas and research topics, such as
Population and Community ecology, Natural history (from a quantitative approach), Infectious disease, and Ecophysiology.
Implications for Conservation: The scientific information gaps represent a challenge in generating effective strategies to
conserve Páramo amphibians, considering the high degree of endemism and threats to these species. More than 80% of the
Páramo amphibian species only have the information of their descriptions and little is known about their ecological re-
quirements, population size, or data related to specific threats.
Keywords
anura, caudata, high mountain, knowledge shortfalls, neotropics, publications, páramo ecosystem
Introduction
The Neotropical high montane includes a variety of unique 1
Pontificia Universidad Javeriana, Facultad de Estudios Ambientales y Rurales.
ecosystems on the planet, including montane and high An-
Carrera 7 N 40 – 62, Bogotá, Colombia.
dean forests and large areas dominated by native grasses 2
Pontificia Universidad Javeriana, Facultad de Estudios Ambientales y Rurales,
(Gradstein et al., 2008; Hofstede, 2013). The tropical alpine Departamento de Ecologı́a y Territorio. Carrera 7 N 40 – 62, Bogotá,
grasslands of America are located on mountain peaks at Colombia.
elevations above 3,000 m. The humid and dry punas of the
Corresponding Author:
Andes, the high-altitude grassland in southeastern Brazil, and Liliana Patricia Saboyá Acosta, Pontificia Universidad Javeriana, Facultad de
the equatorial Páramos are classified within these grasslands Estudios Ambientales y Rurales. Carrera 7 N 40 – 62, Bogotá, Colombia.
(Ruiz et al., 2008; Christmann & Oliveras, 2020). Páramos Email: [email protected]
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2 Tropical Conservation Science
are part of the high montane ecosystems with the largest approach quantify and describe patterns in scientific publica-
extension, being distributed in the equatorial Andes and tions and identify information gaps (Aldana-Domı́nguez et al.,
southern Central America, above the limit of the high Andean 2017; Arbeláez-Cortés, 2013a; Urbina-Cardona et al., 2023).
forests and below the snow line (Lauer, 1981; Hofstede et al., There are some bibliometric analyzes in Colombia about birds
2003; Anthelme & Peyre, 2019). (Estela et al., 2010), turtles (Bock & Páez, 2017), herpetofauna
In Colombia, Páramos are considered strategic ecosystems (Urbina-Cardona et al., 2023), and tropical dry forest ecosystem
of great importance due to the ecosystem services they (Aldana-Domı́nguez et al., 2017), as well as a review on national
provide, such as carbon sequestration and the regulation of biodiversity (Arbeláez-Cortés, 2013b). Bernal & Lynch (2008)
almost 70% of the country’s drinking water (Rivera Ospina & reviewed the geographic distribution and altitude ranges of
Rodrı́guez, 2011; Hofstede, 2013; Cabrera, 2014). The Co- amphibian species in the Colombian Andes, but the state of
lombian Páramos have a unique diversity in terms of species knowledge of the amphibians that inhabit the Páramo ecosystem
richness and endemicity on the planet. This is explained mainly is unknown to date. Systematic reviews are currently popular
by the complex geological and environmental history of the and their important contribution to knowledge is increasingly
Andes (Navas, 1999; Gregory-Wodzicki, 2000; Doan, 2003; recognized, which is related to the high quality and transparency
Madriñán et al., 2013). The high speciation rates are associated that these works have (Siddaway et al., 2019). The aims of this
with differences in environmental conditions within mountain study are 1) to describe the temporal patterns in scientific
systems, which are compared to continental islands productivity between scientific articles in indexed and non-
(Vuilleumier, 1970; Navas, 2002; Mendoza et al., 2015). indexed journals; 2) to characterize the scientific productivity
Amphibians are one of the most diverse lineages of according to the taxonomic groups and the main research topics
vertebrates in Colombian Páramos (Ardila-Robayo & over the years; 3) to identify spatial patterns in the frequency of
Acosta-Galvis, 2000; Lynch & Suárez, 2002). Given their life publications by sector, department, and Páramo complex; 4) to
history and physiological traits, both salamanders and an- evaluate changes in the composition and richness of amphibian
urans have adapted and diversified in these environments. species in the Páramo complexes by sector in Colombia; and 5)
Their low metabolisms rates, behavior, morphological traits, discuss knowledge shortfalls that become challenges for the
and thermal adaptations contributed to compensate for conservation of Páramo amphibians.
overexposure to wind and UV rays or heat loss in an eco-
system where the daily circadian cycle changes abruptly.
Methods
(Navas, 1996a; Navas, 2006; Carvajalino-Fernández et al.,
2011; Navas et al., 2013). Most studies of Páramo amphibians Study area: Páramos are present in Colombia, Costa Rica,
focus on descriptions or species lists (Rueda-Almonacid & Ecuador, Perú, and Venezuela (Hofstede et al., 2003; Cortés-
Hoyos, 1991; Ardila-Robayo & Acosta-Galvis, 2000; Lynch, Duque et al., 2013). In the case of Colombia, this ecosystem
2001; Buitrago-González et al., 2016; Henao et al., 2019), as covers a total of 2.906.137 ha, which is close to 2.5% of the
well as some research in Ecophysiology (Carvajalino- country’s mainland and 50% of the world’s Páramos (Morales
Fernández et al., 2011; Guarnizo & Cannatella, 2014; et al., 2007). Páramos have political and administrative
Navas, 1996b, 1996), morphology (Hoyos et al., 2015; presence in 22 departments, and they are geographically
González-Durán et al., 2017; Mendoza - Henao et al., 2019), present in the Central, Eastern, and Western Ranges, as well
bioacoustics (Fandiño et al., 1997; Gutiérrez & Lüddecke as in the Sierra Nevada de Santa Marta Mountain range
2002) and community ecology (Gutiérrez-Lamus et al., 2004; (SNSM). As an environmental protection policy for the
Roach et al., 2020; Duarte-Ballesteros et al., 2021). However, country, the government created a delimitation process that
despite a large amount of research carried out in the Co- groups most of the Colombian Páramos under the figure
lombian Páramos, scientific publications are scattered in “Páramo complexes” (Morales et al., 2007). This process
various sources and some of them are difficult to access. resulted in a delimitation of 36 complexes grouped into five
The study of anurans and salamanders diversity in the high sectors (Rivera Ospina & Rodrı́guez, 2011; Sarmiento &
mountains of Colombia is of vital importance given the Ungar, 2014; Sarmiento & León, 2015; Figure 1).
accelerated loss of ecosystems in the tropical Andes (Etter & van The Páramo ecosystem is part of the Colombian high
Wyngaarden, 2000; Etter et al., 2008; Etter et al., 2018). Fur- mountains, the geographic space corresponding to the altitu-
thermore, habitat loss in synergy with climate change will in- dinal culminations of the mountainous formations that occur
crease the risk of extinction of amphibian species (Agudelo-Hz from 2700 ± 100 m (Sarmiento et al., 2013). Until now, there is
et al., 2019). These conservation problems are even more ev- no consensus about the altitudinal limits between the eco-
ident in the Páramo ecosystem because its transformation by systems of the Andean, high Andean, and Páramo orobiomes
human actions has recently increased (Alarcón et al., 2002; since the species replacement between these ecosystems varies
Rivera Ospina & Rodrı́guez, 2011; Cadena-Vargas & Sar- greatly throughout their distribution according to their location
miento, 2015; Sarmiento et al., 2017). It is urgent to have a and climatic characteristics (Cuatrecasas, 1958; Lauer, 1981;
database that enables the consolidation of the current knowledge Rangel & Rangel -CH, 2000; Rodrı́guez et al., 2006; Cortés-
of amphibian studies in the Páramo and, from a bibliometric Duque et al., 2013; Hofstede et al., 2014).
Saboyá Acosta and Urbina-Cardona 3
Figure 1. Study area. (A) Distribution of Páramos in the neotropics. (B) Distribution of the 5 sectors within which the 36 Páramo complexes
are grouped, according to the delimitation process of Páramo complexes for Colombia (IAvH, 2012).
Literature search and databases: Our systematic literature Heritage library (BHL) (Biodiversity Heritage, 2021) until
survey included seven different search strategies to detect the 2021. 4) A deep search on the digital repositories of the
largest number of documents published in indexed journals Instituto de Investigación de Recursos Biológicos Alexander
and local sources with low accessibility, between the years von Humboldt - IAvH (2019) (until 2021) and the book series
1863 and 2021: 1) A comprehensive Title-Abstract- “Colombia Diversidad Biótica” (Rangel-Ch, 2019). 5) We
Keywords search in Web of Science (WoS including all consolidate a taxonomic list of Páramo species reported in the
databases: the Core collection, Korean journals and Scielo scientific literature (Acosta-Galvis, 2000, Ardila-Robayo &
citation index) and SCOPUS on the 30th of November 2021, Acosta-Galvis, 2000; Lynch & Suárez, 2002; Bernal &
between the years 1927 and 2021 and using the following Lynch, 2008; Acosta-Galvis, 2015; Buitrago-González
search syntax: (Páramo* OR "andean forest*" OR "Tropical et al., 2016; Meza-Joya & Torres, 2016). 6) We compi-
High mountain" ) AND (anura* OR amphibian* OR tadpole* lated the available records of amphibians present in the
OR frog* OR toad* OR salamander OR caudata OR Biodiversity Information System [by its Spanish acronym
gymnophion* OR caecil *). 2) A search in all the published SiB], and the Global Biodiversity Information Facility
volumes and issues of all Colombian journals indexed by (GBIF), from the delimitation of Páramos of Colombia
MinCiencias (Ministerio de Ciencia, Tecnologı́a e project and the biological collections of the University of
Innovación de Colombia, 2018) with the research topics of Antioquia (MHUA), National University of Colombia (ICN),
natural sciences or biological sciences (online Appendix 1). Universidad del Valle (UVC), Universidad del Quindı́o
3) The same search syntax from search strategy 1 was used (ARUQ) and the Alexander von Humboldt Research Institute
together with the term “Colombia” in the Biodiversity (IAVH) to complement the geographic and taxonomic
Figure 2. PRISMA methodology (Liberati et al., 2009) used in the present study. Acronyms: BHL = Biodiversity Heritage library; IAVH =
Alexander von Humboldt Research Institute; IUCN = International Union for Conservation of Nature; GBIF = Global Biodiversity
Information Facility; SIB = Biodiversity Information System of Colombia
information of the species. And, finally, 7) We searched the documents met at least one of the following inclusion criteria:
bibliographic references for each species (until May 2021) 1) It includes studies in ecosystems above 2700 m in Co-
reported in the International Union for Conservation of lombia; 2) It presents data on species reported in the taxonomic
Nature (IUCN) red list assessments (IUCN, 2021) and the list list compiled from the scientific literature (search strategy 5),
of amphibians of Colombia by Acosta-Galvis (2021). and available records (search strategy 6); and 3) Study areas
We generated a database and manually removed duplicates coincide spatially with some Páramo complex. After a com-
for a total of 882 documents. For each of these documents, the plete review and exclusion of documents that did not meet the
abstract, methods, study area, results, and supplementary selection criteria (n = 477 documents), we consolidated a final
material were read following the PRISMA protocol (Liberati database (Figure 2) comprising 405 eligible documents (online
et al., 2009; Shamseer et al., 2015), to ensure that all eligible Appendix 2) to search the following information:
1. General metadata: Type of document (research article, similarity profiles (SIMPROF routine; Clarke et al., 2008),
book, book chapter, herpetological note, literature review, the grouping between 21 research topics was statistically
data paper, IUCN Red list assessment, SiB-GBIF occurrence validated to identify sets of topics that had similar produc-
data, and Colombian Association of Herpetology catalog files tivity patterns over time.
[by its Spanish acronym CARC], keywords, title, publication, To identify the spatial patterns in productivity, we gen-
year, and source of information: Database: (GeneBank, SiB- erated a database from 1) Each record by species and geo-
GBIF, IUCN), field data, laboratory data, biological collec- graphical unit (department, Páramo complex, sector),
tions, and literature. reported in the documents of online Appendix 2; 2) The
2. Research topics: Adapting the proposal of Urbina- information on the Extent of Occurrence (EOO) from each
Cardona et al., (2023), the first author LPS classified each species of the taxonomic list (search strategy 5) and available
of these documents within some of the following 21 research in the last Red list assessments (IUCN, 2021); and 3) The
topics: Anatomy and Morphology, Bioacoustics, Biogeog- records downloaded from the GBIF and the SIB. For this
raphy, Climate change, Community ecology, Conservation, procedure, we used the “intersection” geoprocessing tool in
Diet, Ecophysiology, Ethology, Evolution, Functional ecol- QGis 3.24.1 (Qgis.org, 2022) and both the occurrences
ogy, Genetics, Geographic distribution, Infectious diseases, (GBIF and SIB) and the EOO in the Redlist (IUCN, 2021)
Molecular biology, Natural history, Phylogenetics, Pop- were extracted and overlapped with the cartographic infor-
ulation ecology, Reproductive biology, Systematics and mation on the Páramo complexes delimitation of Colombia at
Taxonomy, Taxonomic list. A document could be classified a scale of 1:100,000 (IAvH 2012). We excluded from this
within more than one research topic. analysis all the papers related to taxonomic lists that only
3. Geographic distribution: Information on the sector presented general information at the department level without
(Central, Eastern, Western Ranges, Sierra Nevada de Santa reporting records in the Páramo (n = 8). We also excluded 372
Marta -SNSM, and Nariño-Putumayo; See Figure 1), as well records from the literature with uncertain geographic infor-
as on the department and on the Páramo complex (Sarmiento mation about the association of the amphibian species to the
& Ungar, 2014). Páramo ecosystem and 775 GBIF records that did not in-
4. Taxonomic information: The taxonomic list of studied tersect with Páramo complexes.
species, as well as the order and the family. We updated the The final database (online Appendix 3) contains a total of
entire list according to Frost (2021) until January 2022. 2835 records with the following information: species, pub-
Data analysis: Knowledge regarding Páramos amphibians lication, year, Páramo complex, sector, and department.
was explored through descriptive graphs to visualize tem- Given that, in the same study, a species could occur in dif-
poral patterns in publications over the years and by type of ferent departments or Páramo complexes, the information is
document. We used bar plots to contrast the number of papers reported for each of the geographical units (Páramo complex
in scientific journals within the Journal Citation Reports within a sector and in a department). From the online
(JCR) (Web of Science Group - Clarivate) and Scimago (SJR) Appendix 4, the number of publications per geographic
(Scopus) with publications in non-indexed sources to contrast unit was estimated and maps were generated using the QGis
in which source of information (indexed or non-indexed) the 3.24.1 program (Qgis.org, 2022). Similarly, heat maps were
literature of amphibians Páramos is being generated and their created (Somerfield & Clarke, 2013) to explore the associ-
temporal trends. In this way, it can be evidenced if the lit- ation between 21 research topics by departments. And the
erature produced on this topic in the country is remaining in grouping (SIMPROF routine) between research topics and
easily accessible repositories and if they are published in departments was statistically validated in the PRIMER 7.0.13
high-impact scientific journals. Documents were character- and PERMANOVA add on program (Clarke & Gorlye, 2006,
ized according to the 21 research topics for each taxonomic Anderson et al., 2008; Clarke & Gorley, 2015).
order and visualized through a Foam tree (Carrot Search Finally, based on the specific records (museum specimen
FoamTree, 2019). In the Foam tree, each category is divided with data georeferenced or verifiable Páramo locality or EOO
into polygons of different colors and sizes according to their polygon coincidence with Páramo complex; n = 1979), a
class and frequency, respectively. taxonomic list of amphibians was generated for the Colombian
We extracted the information on “Research topics” from Páramo complexes. We excluded from this analysis all
online Appendix 2 to describe the number of publications by specimens that were reported as “sp.” “cf.” or “aff.” or within
research topic and year range and we transformed this data species complexes (eg Atelopus “ignescens”) (n = 1101). We run
into a square root to calculate the association index of a multivariate analysis of variance based on permutations
Whittaker (1952). With the Whittaker matrix, we created a (PERMANOVA; Anderson, 2001) to determine the effect of
heat map to show the temporal patterns in the 21 research the geographic sector of the Páramo complex on amphibian
topics. In the heat map, the frequency of research topics is alpha and beta diversities. The Jaccard similarity index was
represented by a color gradient that goes from low association calculated to evaluate changes in the composition of the as-
(cold tones) to a high degree of association (warm tones) semblages, and for richness, the Euclidean distances between
(Somerfield & Clarke 2013). Additionally, based on the the Páramo complexes were also calculated. The analysis was
Figure 3. Temporal pattern of scientific productivity on amphibians from the Colombian Páramo differentiated by the type of document.
performed under a partial sum of squares (type III) and 9999 other hand, 285 documents are not indexed, of which 25.26%
permutations of the residuals under a reduced model. When correspond to articles published in scientific journals, 46.26%
differences between sectors were found, a student’s t-test to evaluations of the IUCN Red list assessments, 12.45% to
was calculated as a post hoc test. Finally, we explore the book chapters, 8.54% to occurrences from SiB-GBIF, 4.27% to
association of the presence of species by Páramo complex the catalog of the CARC catalog files, and 3.20 % to books.
and sector with a heat map (Somerfield & Clarke, 2013). The The 43.75% of the non-indexed publications are Colombian
classification of species was validated with the SIMPROF and within these, we found two scientific journals (Revista de
routine in PRIMER 7.0.13 and PERMANOVA add on la Academia Colombiana de Ciencias Exactas, Fı́sicas y
program (Anderson et al., 2008; Clarke & Gorley, 2015). Naturales and Biota Colombiana) and two books: El Libro
Rojo de Anfibios de Colombia (Rueda-Almonacid et al., 2004)
and Ranas Arlequines (Rueda-Almonacid et al., 2005) as the
Results most representative from this group (Figure 4B).
Scientific productivity on Páramo amphibians: Types
of publication and temporal trends Taxonomic and thematic characterization of
scientific literature
The 405 documents were published between 1863-2021 and
included: 160 research articles, 134 IUCN species Red list A total of 142 species were recorded within the Amphibia
assessments, 35 book chapters, 29 SiB and GBIF data oc- class with distribution in Colombia’s Páramo complexes:
currence reports, 15 herpetological notes, 12 CARC catalog Order Anura with 134 species in six families, and Order
files, 8 review articles, 9 books, and 2 data papers. The Caudata with five species in one family (Plethodontidae) (See
number of publications increased by 77.18% since the year online Appendix 4). 95 species of our proposed list are re-
2000, mainly with contributions from the IUCN Red list ported as endemic to Colombia. Most of the publications
assessments, research articles in scientific journals (indexed were about anurans (94.68%), while 20 documents were
or non-indexed), and herpetological notes (Figure 3). reported for salamanders, and no information was found for
The scientific articles (data papers, research articles, re- the order Caecilia in the Páramos of Colombia. At the family
views, and herpetological notes n = 185), were published in 73 level, Strabomantidae presented the highest number of
journals. 115 of these publications (62%) are found in 42 studies (n = 219), of which the genera Pristimantis (n = 201)
journals indexed in the JCR of Web of Science (45.36%) or the and Niceforonia (n= 34) were the most representative. The
Scopus SRJ (54.63%), of which the ones with the greatest second-best represented family in the documents was Bu-
contribution were, in descending order: Caldasia, Zootaxa, fonidae (n = 116) with the genera Atelopus (n = 89), and
Herpetologica and Journal of Herpetology (Figure 4A). On the Osornophryne (n= 32), followed by the family Hylidae
Figure 4. The number of publications on Páramo amphibians in Colombia in (A) journals indexed in SRJ and JRC. (*) correspond to
internationally indexed journals from Colombian and (B) in non-indexed publications.
(n=79) with the genera Dendropsophus (n= 62) and Hylo- (Peters, 1863) (n=44), Hyloxalus subpunctatus (Cope, 1899)
scirtus (n=44). The species with the highest frequency of (n=29) and Atelopus carrikeri Ruthven 1916 (n=27).
appearance in the database publications were Dendropsophus The most frequent research topics within the anuran
molitor (Schmidt, 1857) (n=59), Pristimantis bogotensis documents were Natural history (242), Conservation (138),
Figure 5. Polygons´ visualization of the research topics within which the scientific literature on the amphibians from the Colombian Páramo
separated by taxonomic order was classified.
Figure 6. Heat map of the study research topics on amphibians from the Colombian Páramo over time. The tree on the left shows the
classification of the 21 research topics into 4 groups according to their degree of association (Whittaker index) with the year ranges.
and Systematics and Taxonomy (109). In contrast, the order of which 47,6% were classified in the same group, and the
Caudata was included more often in Taxonomic lists (12), other three groups presented between 1 and 7 research topics
Natural history (10), and Systematics and Taxonomy (8) (Figure 6). The first studies in Biogeography, Ecophysiology
studies (Figure 5). and Bioacoustics, as well as an increase in publications on
The publication on Natural history and on Systematics and topics such as Systematics and Taxonomy, and Natural
taxonomy has been maintained over the years, but the state of history, can be traced back to the 70s. Later, in the 90s,
knowledge on Páramo amphibians in Colombia has been publications on Ecophysiology and Bioacoustics continued to
enriched since the 2000s with other 19 research topics. These increase; and, from the 2000s onwards, new topics have
topics were classified into four groups with similar patterns, appeared such as Infectious Diseases, Community ecology,
Figure 7. Geographic patterns the number of documents on amphibians of Colombian Páramos (A) by department (B) Sierra Nevada de
Santa Marta (C) Eastern range sector, (D) Central range sector, (E) Nariño-Putumayo sector, (F) Western range sector.
Reproductive ecology, Diet, and Climate change. In contrast, Cundinamarca) are grouped with many publications on
research topics in Population ecology, and Ethology have Natural history and Conservation (Figure 8). Cundinamarca
shown stagnation or decline. Geographic distribution, Con- represented 85% of the research topics; followed by Anti-
servation, and Natural history are some of the most repre- oquia, Boyacá, Tolima, Caldas, Cauca, Nariño, Quindı́o, and
sentatives of the last decade. Putumayo, which presented between 10 and 16 topics.
Whereas the departments with the lowest number of publi-
cations were Arauca, Caquetá, Huila, Tolima, Cauca, Norte
Spatial patterns from the scientific literature
de Santander, and Santander, which represented between 1
Of the 22 departments with Páramos in Colombia, Cundi- and 2 research topics.
namarca registered the highest number of studies (n= 93),
followed by Antioquia (n=64), Nariño (64), Cauca (n=61), Composition of amphibians in Páramo complexes and
and Boyacá (59) (Figure 7A). The highest number of pub-
lications were reported in the Eastern ranges sector (n=159),
comparison between sectors
in which 17 Páramo complexes are distributed, and the We found differences in the species richness (Pseudo-F=
Central sector (n=129), in which eight Páramos complexes 13.8; p.perm= 0.0001; size effect = 68%) and in the com-
are present (Figure 7C-D). The Páramo complexes that position of the amphibian assemblages (Pseudo-F= 4.15
registered the highest number of studies were Chingaza p.perm= 0.0001; size effect = 34.3%) between sectors. The
(N=70), in the Eastern range sector; Los Nevados (n=62), in general pattern in assemblage composition shows that Sierra
the Central range sector; and La Cocha-Patascoy (n=56), in Nevada de Santa Marta (SNSM), Cordillera Central, and
the Nariño-Putumayo sector. Nariño-Putumayo presented noticeable differences from
From the number of publications by research topics, the other sectors such as Eastern ranges and Western, which is
departments were classified into nine groups according to the associated with the high rate of species turnover that occurs in
degree of association with research topics. Besides, the 21 the high Colombian mountain. The 142 species were clas-
research topics were classified into 4 groups with similar sified into 42 groups according to their distribution in the 35
patterns showing great thematic diversity between depart- Páramos complexes (Picachos was not included because we
ments. Three of these departments (Nariño, Cauca, and do not report species in this complex) and within the five
Figure 8. Heat map of research topics by Department. The tree on the left shows the classification into 4 groups of the 21 research topics by
their degree of association (Whittaker index) with the departments. The classification of the 22 departments into 9 groups is also evident.
Classification trees represent, with red nodes, those entities that have similar publishing patterns. The degree of association is shown as a
gradient of colors ranging from warm tones (high degree of association) to cold tones (low degree of association).
sectors (Figure 9, Table 1). A high degree of endemicity can collections (Ardila-Robayo & Acosta-Galvis, 2000; Bernal &
be observed in Páramos complexes such as SNSM in which Lynch, 2008). However, our study, revels that a low per-
all species are unique to this area including the genus Ser- centage of this literature is published in scientific journals
ranobatrachus (Arroyo et al., 2022). Complexes like Perijá, indexed in the JCR of Web of Science or the SRJ of Scopus.
located in the north region of the eastern sector, also report The highest percentage of the publications in our database are
endemic species like Pristimantis reclusus (Lynch, 2003) and unindexed and are represented mainly by the IUCN red list
Tachiramantis cuentasi (Lynch, 2003), and the Duende assessments, which are open access (IUCN, 2021). Most
complex (Western Sector) presents unique fauna like Boli- discontinued journals (national or foreign) like Lozania,
toglossa hiemalis Lynch 2001, Pristimantis duende (Lynch, Trianea, Life Science Occasional paper, and Miscellaneous
2001) and Pristimantis xeniolum (Lynch, 2001). Almost all publications (University of Michigan. Museum of Zoology)
the Páramos complexes of the eastern sector share species are found in repositories that are not always easy to track or
with wide distribution in the eastern range like Pristimantis access and do not have major visibility.
elegans (Peters, 1863), P. bogotensis, D. molitor or Bolito- In the present study, we included two documents con-
glossa adspersa (Peters, 1863). On the other hand, complexes sidered grey literature, specifically books and reports from
in the Central sector located to the south (Guancas-Puracé- government institutions. These documents were accessed
Coconucos and Sotará) share around 12 species with the through search strategies 4 and 7, followed in the method-
complexes Doña Juana Chimayoy and La Cocha Patascoy ology. Several authors recognize the importance of grey
(Nariño-Putumayo sector). literature in creating an overview of knowledge for a par-
ticular topic, as noted by Mahood et al. (2013), Hortal et al.
(2015), and Adams et al. (2017). This type of literature often
Discussion
contains anecdotal scientific data or data obtained through an
Based on the 405 published documents, this review is the absent or lax research design (e.g.Christie et al., 2020, 2021).
study with the highest bibliographic, geographical, thematic, Still, it complements state-of-the-art on underexplored sites
and taxonomic coverage of amphibians of the Páramo eco- or understudied species. This is especially relevant to com-
system. Generally, the literature compiling data on Páramo plement conservation actions for populations of native spe-
amphibians was limited to species lists constructed from cies and their habitat, as such measures depend on baseline
literature reviews in scientific journals (Lynch & Suárez, information about natural history, life histories, geographic
2002; Buitrago-González et al., 2016) and in some cases, distribution, and their short-, medium- and long-term spatial
it was complemented with information from biological dynamics. For species with a distribution restricted to
Figure 9. Heat map of species by Páramo complex. The tree on the left shows the classification into 42 groups of 142 species. The
classification trees validated with the SIMPROF routine, represent with red nodes, those entities that have similar distribution patterns by
Páramo complex (see Table 1) and by sector: .= Central; = Nariño-Putumayo; = Oriental; = Sierra Nevada de Santa Marta; and
= Western.
Table 1. Number of Studies and Species Reported by the Scientific Literature in the Páramo Complexes Within Five Sectors of Colombia.
Species restricted
Species / endemic to
Sector Páramo Complex Departments Studies reported these areas
Central range Belmira (1), Chilı́ – Barragán (2), Guanacas – Antioquia, Quindı́o, Cauca, Huila, 129 62 37
Puracé – Coconucos (3), Las Hermosas (4), Risaralda, Tolima, Valle del Cauca
Los Nevados (5), Nevado del Huila – Moras
(6), Sonsón (7), Sotará (8).
Nariño – Chiles – Cumbal (9), Doña Juana – Chimayoy Cauca, Nariño, Putumayo 67 39 24
Putumayo (10), La Cocha – Patascoy (11)
Eastern range Almorzadero (12), Altiplano Cundiboyacense Boyacá, Caquetá, Cesar, Huila, 159 48 42
(13), Chingaza (14), Cruz Verde-Sumapaz Cundinamarca, Norte de
(15), Guantiva – La Rusia (16), Guerrero Santander, Santander, Casanare,
(17), Iguaque – Merchán (18), Arauca, Meta
Jurisdicciones – Santurbán – Berlı́n (19),
Los Picachos, Miraflores (20), Perijá (21),
Pisba (22), Rabanal y Rı́o Bogotá (23), Sierra
Nevada del Cocuy (24), Tamá (25), Tota –
Bijagual – Mamapacha (26), Yariguı́es (27).
Sierra Nevada Sierra Nevada de Santa Marta (28) La Guajira, Magdalena, Cesar 35 9 9
de Santa
Marta
Western Cerro Plateado (29), Citará (30), El Duende Antioquia, Valle del Cauca, Chocó, 43 25 9
Range (31), Farallones de Cali (32), Frontino- Cauca, Risaralda
Urrao (33), Paramillo (34), Tatamá (35)
naturally fragmented ecosystems (e.g., páramo amphibians), Ravelo & Martı́nez, 2019). These works sought to increase
grey literature repositories may represent the only informa- the knowledge of the Páramos fauna and flora. However, the
tion available to guide management and conservation actions. information is not available for all paramos complexes and
Open data provided by databases and reported in our study the reports accessible in the GBIF had more than 30% of
(e.g., GeneBank, IUCN, SiB-GBIF) have accelerated re- taxonomic gaps most related to the Pristimantis genus. Other
search in the field of Conservation biology, as well as in data from these reports just provided “Human Observation”
research topics in Evolutionary biology and Systematics but not a specimen deposited in a scientific collection. Some
(Lacher et al., 2012; Baxevanis & Bateman, 2015). However, factors that could explain the low number of fieldwork are 1)
despite being in an open science context, we found just two the difficult access to these sites and the abrupt conditions of
data papers in our database (Henao et al., 2019; Mendoza- climate and topography, 2) political-economic deficiencies
Henao et al., 2019). Data papers are a type of recent academic (Economic resources, research permits), and 3) public safety
publication that offers relevant information regarding the problems (Arbeláez-Cortés, 2013a). This represents a Lin-
visibility and re-use of data that can be available for public nean shortfall (Hortal et al. 2015) in which there is a large gap
use and maintains both scientific rigor and public confidence for the currently formally described species to approach the
(Roche et al., 2015; Jiao & Darch, 2020; Machuca-Martı́nez, total number of amphibian species that inhabit the Páramos
2020). This is of great importance as the public availability of and are still unknown to us (Brito 2010).
primary biodiversity data is essential for ecological research
(Huang et al., 2013), future meta-analyses, and decision-
Thematic information gaps
making in critical ecosystems such as the Páramo. However,
the low frequency of these types of papers may be because the From the first study reported in 1863 to the years 2010-2021,
authors 1) do not want to share their primary data before they the generation of new knowledge published in scientific
are published; 2) do not want to undergo a peer review articles is mainly dominated by research topics such as
procedure again, which may place an additional burden; or 3) Systematics and Taxonomy, Natural history, and Anatomy
do not want to subject their data to a statistical reanalysis by and Morphology. Most of the indexed publications covered
other colleagues (Huang et al., 2013; Wicherts et al., 2011). these topics in descriptions of new species or taxonomic
To advance the state of knowledge of Páramo amphibians it is reviews at the family level (Duellman & Hillis, 1987;
essential that authors have the good practice of uploading Duellman et al., 2016; Estupiñan et al., 2016; González-
their raw data in repositories (e.g. Dryad, Mendeley Data or Durán et al., 2017; Rivera-Correa et al., 2017). In many cases,
ResearchGate) and report the download link in their these descriptions encompassed anatomical aspects of species
publications. and life history data. However, in most cases, the latter topic
On the other hand, the biological collections provided data has been covered from anecdotal narratives or qualitative
for 56.45% of the investigations reported in scientific articles descriptions (Coloma et al., 2000; Hedges et al., 2008; Páez-
(Data papers, research articles, reviews, and herpetological Moscoso et al., 2011). In recent decades, the importance of
notes), mainly related to research topics such as Systematics data related to life history has been highlighted since this
and Taxonomy, Anatomy and Morphology, Natural history, information allows a better understanding of the ecological
taxonomic lists, biogeography, and geographical distribution. strategies of species for conservation (Becker et al., 2010;
These data highlight the importance of collections as sources Michaels et al., 2014); as well as their possible response to the
of scientific data to document the country’s diversity (Medina current scenarios of climate change, land use, and land cover
et al., 2016). Nevertheless, the lack of availability and up- change that our planet is experiencing (Oliveira et al., 2017).
dating of records in several of the collections consulted in the In fact, we only found one study on climate change (Agudelo-
GBIF is notorious. The largest number of specimens reported Hz et al., 2019). This is alarming due to the current challenge
in our database are concentrated in the collections of the ICN that Colombia faces in relation to having data to measure the
and IAvH, but many of them lack georeferencing and alti- effects of climate change on appropriate time scales, as well
tudinal data. Other specimens are omitted in these GBIF as modeling and understanding interactions in ecosystems,
reports and the oldest ones (mainly those of the nineteenth both between biotic components and with other external
and twentieth centuries) present errors related to their lo- transformation drivers (Londoño et al., 2019). For this reason,
calities that require an update (Vásquez-Restrepo, 2021). it is important to increase studies on this theme, but these
We report a low percentage of studies whose data source is should include quantitative and population information
from fieldwork (24.19%) or laboratory data (2.22%). The lack considered that Natural history data may be the key to the
of field information contributes to the underestimation of the success of ex-situ conservation programs (Michaels et al.,
amphibian diversity reported for the Páramos. There are 2014).
recent efforts related to explorations promoted by academic From the 90s onwards, there has been an increase in topics
or governmental institutions like Expedition BIO- such as Ecophysiology, Biogeography, Community ecology,
MinCiencia; and Delimitation of Páramos of Colombia and Bioacoustics; but there is still evidence of an information
(Acosta-Galvis & Borja-Acosta, 2021; Henao et al., 2019; gap on themes like Infectious diseases, Functional ecology,
Climate change, Ethology, and Evolution. Not to mention that nominal species but morphologically indistinguishable be-
there are few studies carried out in Population ecology, which tween populations (Bickford et al., 2007). These gaps in
is a deficiency that is evident in herpetology studies in Co- knowledge from the cryptic species increase the Linnean
lombia (Urbina-Cardona et al., 2023). Despite the incorpo- shortfall (Walters et al., 2021) for amphibians inhabiting the
ration of new research topics in recent decades, studies related Páramos of Colombia.
to field or laboratory data collection have only concentrated in Due to the ecological conditions related to the complex
19.58% of the species in our database. Just 17 Páramo history of the Andean Mountain ranges (biogeography, to-
complexes have at least one study different from the de- pography, and glacial geomorphology), the description of
scription of new species, being the complexes of Los Ne- new species in high mountain ecosystems is to be expected.
vados (Central Sector) and Chingaza (Eastern sector) the This is because it is a pattern consistent with the hypotheses
most representative. of allopatric and peripatric speciation in amphibians dis-
The increase of information available for species in re- tributed in the neotropical Andes and particularly in the
search topics such as "geographical distribution" and "con- Páramos, where the assemblages are unique within each
servation"(since the 2000s) is mainly associated with the mountain system and with a high degree of endemism (Lynch
update made by the IUCN with the Red List assessments & Suárez, 2002; Bernal & Lynch, 2008; Santos et al., 2009;
between the years 2016 – 2021 and the publication of the Garcı́a-R et al., 2012; Mendoza et al., 2015). Hence the
Colombian Association of Herpetology catalog (CARC) files, importance of filling these shortfalls since the prioritization of
that compiles information on taxonomy, life history, mor- conservation efforts at the regional scale is usually deter-
phology and geographic distribution of the Colombian am- mined by the richness of species as a unit of analysis and their
phibian and reptile species; the CARC also provide endemism (Riddler & Hafner, 1999; Fleishman et al., 2006;
information on threats and conservation status. These doc- Mendoza et al., 2015, Espı́ndola et al., 2016). The identifi-
uments have been key to collecting secondary and biological cation of cryptic diversity or the description of new species
collection information, not only on Páramo amphibian spe- from an integrative taxonomy (phylogeography, comparative
cies but also on the herpetofauna of Colombia (Urbina- morphology, population genetics, ecology) and fieldwork in
Cardona et al., 2023). unexplored Páramos is essential to estimate these parameters
in the Colombian high mountains. In addition, taxonomic
work has a strong impact not only on the generation of new
Taxonomic information gaps
knowledge of species, but also on international agreements
Colombia has the largest extension of Páramo in the world on biodiversity conservation, environmental legislation, and
and the highest richness of amphibian species (142) in this distribution of economic resources that are allocated to the
ecosystem, compared to countries such as Costa Rica (n=14; species monitoring programs (Garnett & Christidis, 2017; de
Kappelle & Savage, 2005), Ecuador (n=56; Ron et al., 2020), Magalhães et al., 2018; Thompson et al., 2018). Therefore, it
Peru (n=35; Catenazzi & Von May, 2014) and Venezuela is important not to take for granted the implications of
(n=14; Barrio-Amorós, 2004). However, the diversity of splitting species. Moreover, for the Páramo ecosystem, it is
amphibians reported in Colombian Páramos is yet to be necessary to increase efforts in genuine taxonomic reviews
completed and it requires a greater sampling effort and genetic and not only provide simple lists that end up being taken by
analysis (Ardila-Robayo & Acosta-Galvis, 2000; Lynch & environmental authorities as databases for decision-making.
Suárez, 2002; Acosta-Galvis, 2015). Our database reports a Furthermore, we found few records (<2) or uncertain
significant number of specimens that were cataloged, in records (unverifiable locations) of species that share distri-
scientific articles (>100) and data from the SiB-GBIF (>600), bution in adjacent countries. In our database, we highlight the
as “sp.” and in some cases with annotation of their future case of species contiguous to Ecuador such as Pristimantis
description (Buitrago-González et al., 2016; Acevedo et al., calcarulatus (Lynch, 1976), Pristimantis ortizi (Guayasamin
2018; González-Durán et al., 2017), or species reported as et al., 2004), Pristimantis huicundo (Guayasamin et al., 20
“cf.” or “aff” due to uncertainty in their identification 04), or Pristimantis leoni (Lynch, 1976), which only have
(Cisneros-Heredia & Gluesenkamp, 2010; Acevedo et al., occurrences from the SiB-GIF or IUCN for the Nariño Pu-
2018; López, 2017; Carvajal-C et al., 2019). Identifying tumayo sector (Solarte Cruz, 2021). Similarly, towards the
discrete biological units is fundamental not only for taxon- eastern mountain range adjacent to Venezuela, no records
omy but also for understanding the processes that lead to were found for the species Atelopus tamaense (La Marca,
lineage diversification and defining conservation strategies Garcı́a-Pérez, & Renjifo, 1990) in the Tamá Páramo Complex
(Espı́ndola et al., 2016; Theodoridis et al., 2019). The lack of in Colombia. For the species Hyloscirtus platydactylus,
assigning a name to several specimens captured in these (Boulenger, 1905) the IUCN (2021) reports uncertainty of its
expeditions and reported in the SiB-GBIF or scientific articles presence in the Perijá mountain range, but Carvajal-C et al.,
shows that quantifying the diversity of Páramo amphibians (2019) and Moreno-Arias et al. (2007) report this species
could be hindered by the presence of a cryptic diversity, (although not for the Páramo zone). And for the character-
which is defined as the deep genetic divergence within ization of the Perijá Páramo complex, georeferenced
specimens are recorded and deposited in the ICN biological present records of field data or vouchers in biological col-
collection (Borja-Acosta 2022; Cantillo, 2022). lections. Similarly, despite the delimitation of 36 Páramo
complexes in the country, we still report an information gap in
Miraflores, Los Picachos, Yariguı́es, Paramillo, Citará, and
Geographic information gaps
Cerro Plateado, of which no publication was found in sci-
We included 18 species in our list given the coincidence of entific articles, books, or herpetological notes. The infor-
their EOO (IUCN 2016-2021) with some Páramo complex. mation presented for these geographic units corresponds to
Similarly, only 94 species of the reported 142 presented the occurrence data reported in the SiB-GBIF and the EOO
occurrence data with verifiable information in the GBIF (See coincidence of some species reported in the literature as
online Appendix 4). The lack of adequate georeferencing in “amphibian Páramos” and generated by the IUCN (2021).
the studies and in the largest biological collections in the This fact shows the lack of intensive studies in the south of the
country (like ICN and IAvH) before the year 2000 is one of Eastern range and north of the Central and Western ranges
the most critical problems (Vásquez-Restrepo 2021) that (Ardila-Robayo & Acosta-Galvis, 2000). Also, according to
makes it impossible to have certainty of the distribution of Acosta-Galvis (2015), the information provided for most
some species of amphibians in the altitudinal range of the Colombian Páramos cannot be considered as complete
Colombian high mountain (High Andean Forest, sub- characterizations. There are many challenges in the Co-
Páramo, Páramo, or super-Páramo). Such is the case of lombian high mountains, as well as information gaps that
species like Rhinella nicefori (Cochran & Goin, 1970), need to be filled. While there are departments where the
Rhinella macrorhina, (Trueb, 1971), Atelopus subornatus absence of studies on research topics such as Reproductive
(Werner, 1899), Centrolene notosticta (Ruiz-Carranza & biology, Ecophysiology, Population ecology, and Ethology
Lynch, 1991), Hyloxalus pulchellus (Jiménez de la Espada, are notorious; 85% of the research topics and fieldwork was
1875), Pristimantis tubernasus (Rivero, 1984), Pristimantis concentrated in departments such as Cundinamarca, Anti-
susaguae (Rueda-Almonacid, Lynch, & Galvis-Peñuela, oquia, Boyacá, and Tolima.
2003), Serranobatrachus sanctaemartae (Ruthven, 1917),
and Tachiramantis tayrona (Lynch & Ruiz-Carranza, 1985);
which have been considered in several lists of species as
Implications for Conservation
"paramunas" (Ardila-Robayo & Acosta-Galvis 2000; Colombian Páramos register an alarming degree of inter-
Buitrago-González et al., 2016). This gap in knowledge of vention with the replacement of natural cover and only
the geographic distributions of Páramo amphibians is an 58.33% are within the national system of protected areas
example of the Wallacean shortfall (Hortal et al., 2015) that (Gómez et al., 2015, Cortés-Duque et al., 2013). According to
is present in the neotropics and constitutes a challenge for the analyses carried out by the IAvH (Moreno et al., 2020) on
biodiversity conservation (Bini et al., 2006). a national scale, all Páramo complexes report an increase in
Historically, one of the main criteria used for the delim- transformed coverage from 0.6 to 0.9% due to agricultural
itation of Colombian Páramos has been altitude or elevation activities, ranching, and mining (Burbano-Girón et al., 2020).
(Hofstede, 2013; Hofstede, et al., 2003; Sarmiento & León, On a regional scale, the Eastern, Central, and Nariño-
2015). However, given the geomorphological variety of the Putumayo sectors show a pattern of increase in trans-
country, many Páramos are not homogeneous in their limits, formed areas up to 18.3% in the last decades, which could be
such as the azonal Páramos of the Guamuez River (2500 – associated with the “upward movement” or “upward ex-
2800m) in southern Colombia (Barriga et al., 2015; Ravelo & pansion” of the agricultural frontier and the intensification of
Martı́nez, 2019) or Páramos complexes that start from animal production in the sub-Páramo belt, which has been
2700 m (e.g., Perijá) while others have a lower limit of occurring in different parts of the equatorial Andes (Hess,
3000 – 3500 m or more (e.g., Complexes located in the 1990). These data can be worrying since these sectors account
central mountain range). These differences are influenced by for more than 80% of the species reported as endemic to the
diverse factors such as location, soil types and climate (Lauer Páramo or with populations established in these sites, and
1981; Rodrı́guez et al., 2006), and even the type and degree of these are organisms with restricted areas of occupation
human intervention determines the actual structure and compared to other biological groups. Impacts on the natural
functioning of the Páramo (Hofstede & Llambı́, 2020). The cover can lead to habitat conversion for many species of
altitudinal limits reported for species do not provide all the amphibians. They have a long-term negative effect on am-
necessary information to determine the organisms that are phibian populations and increase the probability of direct
endemic to the ecosystem or those that transit in the altitu- mortality of species (Nowakowski et al., 2017; Catenazzi,
dinal gradient from high Andean Forest - Páramo and that 2015). Currently, 66.19% of Páramo amphibians are within
have been able to establish populations in the Páramo eco- some state of threat: Vulnerable (VU), Endangered (EN),
system (Lynch & Suárez, 2002). Critically Endangered (CR); mainly due to 1) Areas of Oc-
Of the 22 departments with the presence of Páramos in cupation (AOO) between 10 and 2000 km2, associated with the
their territories, Arauca, Casanare, and Caquetá did not low geographical records that exist per species and 2) the
increase in threats related to categories such as “Agriculture & ecology, population trends, and even taxonomy (IUCN,
Aquaculture”, “Residential & commercial development”, 2021). The knowledge shortfalls identified in this paper
“Invasive and other problematic species, genes & diseases” and (Linnean, Wallacean, Prestonian shortfalls; Hortal et al.,
“Energy production & mining” (Stuart et al., 2008, IUCN, 2015) show thematic, spatial, and temporal biases that
2021). skew the possible evidence on the conservation (sensu
The trend towards the transformation and replacement of Christie et al., 2020, 2021)) of amphibians inhabiting the
natural Páramo cover by crops or grazing areas shows that páramos of Colombia. In a regional context, this is of great
habitat conservation is a high priority for the persistence of importance since the tropical Andes is the place in the world
amphibian species. Studies carried out in some Páramos show with the largest number of amphibian species with protection
the use of microhabitats associated with endemic plants from figures gaps in terms of protected areas (Nori et al., 2015).
these areas, such as cushions and shrubs, tussock grasses These data reveal the need to increase the knowledge not
(Calamagrostis spp.), rosette plants and stemmed rosette only of the Páramos inventory but of the biology and dis-
plants of Espeletia spp (Lynch, 1998; Lynch, 2000; Acosta- tribution of the species that inhabit them, because this in-
Galvis, 2015). Also, it is important to note that, at finer spatial formation contributes to the generation of monitoring
scales, each Páramo can generate a unique variety of mi- programs and conservation actions focused on the attributes
croclimates or microecosystems to which amphibians could of the habitat of the species. Moreover, the information
be adapted (Navas, 1996a; Buitrago-González et al., 2016; generated in the scientific literature and records in databases
Duarte-Ballesteros et al., 2021). However, the absence of (such as SiB and GBIF) can strengthen other initiatives like
population data or quantitative records of natural history data the Colombian community of practice called "Biomodelos"
for most species is notorious, without regard to the large gaps (http://biomodelos.humboldt.org.co/es), which seek to gen-
that exist on a spatial and thematic scale and the concentration erate more accurate distribution models and in which only 30
of studies on a few research topics, complexes of Páramos, species (only 21.12%) of our list are reported. Once validated
and even common species. The gap in knowledge of species by experts, the distribution of species in Biomodelos is used
population dynamics is called a Prestonian shortfall and as input to calculate different metrics including AOO and
represents a challenge from long-term monitoring to as- EOO, distribution in protected areas, and changes in distri-
sessing the risk of species extinction and implementing bution under future scenarios of landscape transformation
management and conservation actions for populations and (Velásquez-Tibatá et al., 2019). For this reason, it becomes an
their habitats (Hortal et al., 2015). More than 80% of the indispensable tool to make the processes of amphibian
species that inhabit or transit the Páramo in Colombia only extinction risk assessment in Colombia more efficient
have their descriptions information or have been mentioned (Urbina-Cardona et al., 2023). It is important to highlight that
in the literature in species lists but little is known about their generating data in situ involves a greater sampling effort, the
ecological requirements or data related to specific threats: use of specialized equipment, and reagents for tissue col-
Infectious diseases, invasive species, and climate change. lection or animal experimentation. These are activities that
On the other hand, current information gaps related to the require greater financial support, which is a challenge since
assignment of the nominal status of species and errors in the country is currently investing > 1% of its gross domestic
geographical records can affect categorization in Red Lists, as product in science, technology, and innovation (The World
they cause inaccuracy in the estimation of the EOO and AOO. Bank, 2018; Guevara, 2021).
Consequently, we could be hiding the status of some species
with degrees of threat (de Magalhães et al., 2018). Currently, Acknowledgments
eight of the species with a presence in the Páramo are found We thank Ana Marı́a Suárez for assistance with Maps in QGis,
with Deficient Data (DD). Two of them are uncategorized, Camila Rodrı́guez for her comments on English version and Pon-
and 80.85% of the species endemic to certain Páramos tificia Universidad Javeriana for academic database access.This
complexes are distributed only in one “Protected area” manuscript is the fourth publication of the Semillero Javeriano en
(natural parks or private ecological reserves) as a conser- Ecologı́a y Conservación de Anfibios y Reptiles-SECAR.
vation figure (See appendix 4). Although a few studies report
the use of good practice protocols for amphibian sampling Declaration of Conflicting Interests
(e.g., using gloves to handle animals), there is a clear lack of
efforts to mitigate threats to the habitat of these species or to The author(s) declared no potential conflicts of interest with respect
implement concrete actions for their conservation. (i.e., to the research, authorship, and/or publication of this article.
amphibian translocation, citizen science, engagement of
landowners and other volunteers in managing land for am- Funding
phibians, or management plans with the support of envi- The author(s) disclosed receipt of the following financial support for
ronmental institutions; Sutherland et al. 2019. This lack of the research, authorship, and/or publication of this article: This work
effort is mainly due to the absence of field data, including was supported by the Vicerectory of Research, for the support for
population size, distribution and trends, life history and doctoral thesis projects of the Pontificia Universidad Javeriana
(VRI05-19) (Grant ID 20083_120130Z0401200) with the project: Aldana-Domı́nguez, J., Montes, C., Martı́nez, M., Medina, N.,
Morphological and physiological responses of anuran species to Hahn, J., & Duque, D. (2017). Biodiversity and ecosystem
environmental heterogeneity at different scales: the relationship services knowledge in the Colombian Caribbean: progress and
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Saboya Acosta Urbina Cardona 2023 Current State of Knowledge of Paramo Amphibians in Colombia Spatio Temporal Trends

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Saboya Acosta Urbina Cardona 2023 Current State of Knowledge of Paramo Amphibians in Colombia Spatio Temporal Trends

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Tropical Conservation Science

Liliana Patricia Saboyá Acosta1  and J. Nicolás Urbina-Cardona2

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