The rainfrogs of the genus Pristimantis are one of the most diverse groups of vertebrates, with outstanding reproductive modes and strategies driving their success in colonizing new habitats. The rate of Pristimantis species discovered annually has increased continuously during the last 50 years, establishing the remarkable diversity found in this genus. In this paper the specifics of publications describing new species in the group are examined, including authorship, author gender, year, language, journal, scientific collections, and other details. Detailed information on the descriptions of 591 species of Pristimantis published to date (June 2022) were analyzed and extracted. John D. Lynch and William E. Duellman are the most prolific authors, yet Latin American researchers have scaled up and continued the description processes since the 1990s. The most common language used for descriptions is English, followed by Spanish. The great majority of authors have described only one species. The largest proportion of authors who have participated in the descriptions is of Ecuadorian nationality. Ecuador is the country with the highest description rate per year (3.9% growth rate). Only 20% of the contributions have included women and only 2% have featured women as principal authors. 36.8% of the species described are in the Not Evaluated or Data Deficient categories under the IUCN global red list. The importance of enhancing the descriptions in Spanish is emphasized and the inclusion based on equal access to opportunities for female researchers in Pristimantis taxonomy is encouraged. In general, if the current trends in Pristimantis descriptions continue, in ten years, a total of 770 or more species described could be expected.

Las ranas de la lluvia del género Pristimantis es uno de los grupos de vertebrados más diversos, con una variedad de modos reproductivos y estrategias que impulsan su éxito en la colonización de nuevos hábitats. La tasa de especies de Pristimantis descubiertas anualmente ha aumentado continuamente durante los últimos 50 años, estableciendo la notable diversidad encontrada en este género. En este artículo, examinamos los detalles de las publicaciones que describen nuevas especies en el grupo, incluida la autoría, el año, el idioma, la revista, el género, las colecciones científicas y otros detalles. Analizamos y extrajimos información detallada sobre las descripciones de 591 especies de Pristimantis publicadas hasta la fecha (junio 2022). John D. Lynch y William E. Duellman son los autores más prolíficos, pero los investigadores latinoamericanos han ampliado y continuado los procesos de descripción desde la década de 1990. El idioma más común utilizado para las descripciones es el inglés, seguido del español. La gran mayoría de los autores han descrito una sola especie. La mayor proporción de autores que han participado en las descripciones es de nacionalidad ecuatoriana. Ecuador es el país con la tasa de descripción más alta por año (tasa de crecimiento del 3,9%). Solo el 20% de las contribuciones han incluido a mujeres y solo el 2% las ha presentado como autoras principales. El 36,8% de las especies descritas se encuentran en las categorías No evaluadas o Datos insuficientes de la lista roja mundial de la UICN. Destacamos la importancia de potenciar las descripciones en español y fomentar la inclusión de mujeres investigadoras en la taxonomía de Pristimantis. En general, si continúan las tendencias actuales en las descripciones de Pristimantis, en 10 años se podría esperar un total de 770 o más especies descritas.

Author gender, herpetology, inclusion, language bias, new species, taxonomy

Pristimantis Jiménez de la Espada, 1870 is a clade of New-World direct-developing frogs belonging to the family Strabomantidae, order Anura, class Amphibia, phylum Chordata. It is the most speciose genus of terrestrial vertebrates with 591 described species to date (Hedges et al. 2008; Frost 2022). A greater focus on molecular, acoustic, and osteological techniques combined with a significant increase in sampling efforts has led to a rise in the number of newly described species in recent years, allowing further research in understanding their taxonomy and systematics (Padial et al. 2010; Hutter and Guayasamin 2015; Kaiser et al. 2015; González-Durán et al. 2017; Reyes-Puig et al. 2020).

The earliest description of this genus was the description of the genus type Pristimantis galdi (Jiménez de la Espada, 1870). The genus was later placed under the synonymy of Hylodes sensu lato by Boulenger (1882), then synonymized as Eleutherodactylus by Peters (1955). Cyclocephalus, Pseudohyla, and Trachyphrynus were also synonymized with Eleutherodactylus by Lynch (1968, 1971). Heinicke et al. (2007) removed Pristimantis from the synonymy under Eleutherodactylus, with support from molecular evidence. This large and phenotypically diverse genus has faced subsequent molecular analyses confirming its monophyly and status as a closely related taxa to Lynchius, Oreobates, and Phyrnopus, with Yunganastes being suggested as well (Pyron and Wiens 2011; Canedo and Haddad 2012; Pinto-Sánchez et al. 2012). Several discernible groups can be found within the genus, originally described based on extensive morphological data and revised on subsequent molecular analyses (Lynch and Duellman 1997; Hedges et al. 2008; Pinto-Sánchez et al. 2012; Padial et al. 2014; Mendoza et al. 2015). There are currently 13 recognized species groups, with P. conspicillatus as the largest with 36 species (Padial et al. 2014; González-Durán et al. 2017; Reyes-Puig et al. 2020; Taucce et al. 2020). Several other species remain unassigned as they are demonstrably non-monophyletic. These species can maintain taxonomic value as they can be grouped among phenotypically similar species, thus revealing useful comparative information (Hedges et al. 2008; Padial et al. 2014).

Members of this genus are remarkable for laying eggs in terrestrial habitats, with the embryos developing directly into frogs, bypassing the aquatic stage of their lifecycle (Woolbright 1985; Duellman and Lehr 2009). Assemblages of Pristimantis species are common, as their morphology and therefore behavioral and ecological activities are remarkably similar (Arroyo et al. 2008). They are characterized as insectivorous generalists, choosing prey depending on availability and size (Lynch and Duellman 1997; Arroyo et al. 2008). Individuals are predominantly arboreal and nocturnal, commonly perching on leaves at heights below 200 cm. As their reproductive biology leads them away from congregating at ponds, a common strategy for males in this genus is to vocalize from the ground or a suitable perch in search of a mate (Lynch and Duellman 1997; Duellman and Lehr 2009).

The genus is widely distributed throughout the New World and considered the most extensive among Neotropical amphibians, with species found in tropical and subtropical forests in South America and up to lower Central America (Lynch and Duellman 1997; Pinto-Sánchez et al. 2012; Meza-Joya and Torres 2016; Armesto and Señaris 2017). The group shows particularly high levels of diversity and endemism in the Andes, predominantly at elevations above 2000 m along humid environments and surrounding lowlands and Chocoan South America (Heinicke et al. 2007; Meza-Joya and Torres 2016; Armesto and Señaris 2017; Reyes-Puig et al. 2020). The diversity of the genus is greater in Colombia, Ecuador, and Peru and the piedmont, montane, and montane cloud forests of the western and eastern slopes of the three countries concentrate the highest levels of endemism (Hedges et al. 2008; Ron et al. 2022).The continuous increase in the number of species described within this genus suggests a much greater species richness than anticipated, with subtle differences in behavior and physiology reflecting distinct evolutionary pathways (de Queiroz 2005; Hutter and Guayasamin 2015).

The conditions leading to such a high diversification rate in Pristimantis are not completely understood. Families of direct-developing frogs diverged quickly during the early to middle Cenozoic, favoring a wide dispersion across a range of habitats in South America, leading to a rapid accumulation of population genetic isolation (Heinicke et al. 2007; Heinicke et al. 2009; Pröhl et al. 2010). An important point of the radiation of the genus is found between 1000 and 3000 m in the northwestern Andes (Mendoza et al. 2015). The complex biogeographic dynamics in the area not only supported allopatric speciation, but also facilitated dispersion of lowland species during the Paleocene and Pliocene, leading to a speciation pattern particularly suitable to originate cryptic and sibling lineages (Lynch and Duellman 1997; Mendoza et al. 2015).

In the last 20 years, the increase in species description rates in South America has shed light on the remarkable diversity and endemism of Pristimantis, while hinting at the complex patterns of speciation taking place. Given the cryptic nature found in members of this genus, the work to discover and describe new species appears far from over. Categorizing these frogs and their characteristics in a taxonomic context presents several challenges, further obscured by the high degree of plasticity evidenced across the group (Guayasamin et al. 2015).

Despite the fact that this genus is so diverse and taxonomic and systematic research continues from year to year, information on publication patterns and trends is scarce. Issues such as gender and language biases in publications have not yet been explored. Within the biological sciences, Zoology in particular, studies in this branch have been characterized as male-dominated, imposing limitations in the professional development of many women (Slobodian et al. 2021). Similarly, it has been shown that, for example, in ecology and zoology the proportion of principal investigators publishing with women is lower compared to the proportion with men (Salerno et al. 2019). On the other hand, the language of publication continues to be dominated by the English language, although there have been approaches to improve the transmission of science so that it can be disseminated locally (Ramírez-Castañeda 2020). Latin American researchers not only tend to be under pressure to publish in English, but also to do so with colleagues from developed countries, issues that tend to be related to the number of citations an article can receive (Meneghini et al. 2008). Thus, in this article we aim to delve into the specifics of the biases and trends in publication of descriptions of new species of Pristimantis by carrying out a detailed review of all description parameters, emphasizing location, authorship (including author gender), scientific collection, and language.

We followed the proposal of Hedges et al. (2008) and Heinicke et al. (2018) for family classification of Pristimantis. The updated list of species formally described to date was extracted from the Amphibian Species of the World web page from the American Museum of Natural History (Frost 2022). On 14 June 2022, the Pristimantis list contained 591 described species. Based on this list, we generated a detailed database for each species, extracting specific data available on the original description and the aforementioned web page. We built a database with the following fields: unique species identifier, species, journal, year of publication, first author, authors, gender of the author, nationality of authors, corresponding author, genus, country of description, type locality, holotype, synonymy, species group (if applicable), distribution, language of the description, institutional affiliations, conservation status and type of description. For more details see Suppl. material 1. We calculated the growth rate of global descriptions and that of the seven countries with the highest number of descriptions (i.e., Ecuador, Colombia, Peru, Venezuela, Brazil, Panamá, Bolivia). In order to better represent the different historical trends seen in Pristimantis description, growth rates are divided into two periods: The first from 1958–1989, when descriptions begin to be more constant, and the second one from 1990–2021. Due to a significant increase in descriptions at the time, the period ranging from 2010–2021 is also considered. This is mainly because we intend to identify a realistic description rate with the latest advances in the taxonomy and systematics of the genus. In addition to the fact that the number of researchers currently working in Pristimantis taxonomy is greater than in previous periods (Suppl. material 1). We did not consider the periods prior to 1958, as there is no significant growth in description rates observed during them.

We carried out the search for publicly available information regarding gender and nationality of authors through individual exploration enabled by the Google search engine. This search was based on exploration of the Google search results associated with the names of authors (as self-reported in publications describing Pristimantis species) throughout the web. Priority was given to information associated with institutional affiliation and research endeavors, cross-referenced with professional sites such as ResearchGate, LinkedIn and Google Scholar. Information regarding the gender of authors was assessed based on a binary understanding of gender and assumed based on gender roles traditionally assigned to their names, physical appearance, and self-reported identity, when available. Information regarding conservation status was obtained from the IUCN Red List of Threatened Species (IUCN 2022). After finalizing data input, we cleaned and homogenized the database to prevent typographical errors as well as duplicate and empty cells. In order to describe the increase in women involved in Pristimantis descriptions over the years, we used generalized linear models (GLMs). These include a quasi-binomial error distribution, necessary given the nature of the data (i.e., dichotomous proportionality data and overdispersion). We defined years as the dependent variable and the proportion of female authors as the independent variable, in order to identify the relationship between the proportion of women featured in descriptions and time. We made a GLM of the total number of female authors and one specific to each of the five countries with the greatest diversity of Pristimantis (Ecuador, Colombia, Peru, Venezuela, and Brazil). The timeframe for this analysis is from 1970 onwards, corresponding to the period in which women begin to be active in describing a new species of Pristimantis. All management, cleaning, and analysis of the database were performed in the statistical software R (R Core Team 2022); utilizing the “tidyverse”, “forcats” and “gridExtra” packages.

Species descriptions of the group began in 1858 with Pristimantis conspicillatus (under the synonymy of Hylodes conspicillatus, Lithodytes conspicillatus, and later Eleutherodactylus conspicillatus). For a century the descriptions of this group remained relatively stable with fewer than four species described on average every ten years (Fig. 1). From the 1960s, the descriptions in the group increased significantly with several peaks towards the 70s but with a particular one towards 1980. The number of descriptions every five years increased in relation to the range between 1958–1970, with an average of 5–14 descriptions between 1978 and 1996. After this year, the most notable peaks in new descriptions were restricted to the years 1997, 1998, 1999, 2007, 2019, 2020, and 2021 (Fig. 1A). The significant increase in the descriptions of new species in this group intensified after 1978 and the accumulation curve of descriptions seems to continue increasing up to the present date, with a total of 591 formally described species (Fig. 1B). The period with the highest annual rates of Pristimantis description is found between 1958–1989. For Ecuador, the period with the highest description rate lies between 2010–2021, for Colombia between 1958–1989, for Peru between 1990–2021, and for both Brazil and Bolivia, a peak in the description rate is observed in the period 2010–2021 and 1990–2021, respectively. However, it is important to consider that in order to calculate the growth rate, the initial and final values of the numbers of described species are taken into account. By having such a low number of total descriptions, if the species double from one year to the next, the rate is higher; however, the total number of species is much lower compared to that of other countries (Table 1).

Figure 1. 

Pristimantis descriptions through time A species description over 164 years, dots and green lines represent new taxa per year, diamonds and brown lines five-year average, and triangles and purple lines ten-year average B cumulative number of species descriptions.

The distribution of Pristimantis extends from Honduras to Peru, specifically, Honduras east through Central America through Colombia and Ecuador to Peru, Bolivia, northern Argentina, and Amazonian and Atlantic Forest Brazil and the Guianas; Trinidad and Tobago; and Grenada, Lesser Antilles (Frost 2022). Ecuador is the country with the highest number of described species with 212 total descriptions, followed by Colombia (168 descriptions), Peru (107 descriptions), and Venezuela (56 descriptions). Other countries within the range of this genus report fewer than ten descriptions (Fig. 2A). From the 1960s onwards, descriptions occurred predominantly in Ecuador, Colombia, and Peru, with at least two peaks of descriptions higher than the average of ten new species described per year (Fig. 2B). Ecuador exhibits three important peaks in species description (i.e., 1979, 1980, and 2019), with more than 15 new species described per year; Colombia peaks in 1998 reaching 20 descriptions in a single year, setting a record in descriptions for this genus; and finally, Peru peaks twice (i.e., 1999 and 2007) reaching 13 and 15 species descriptions, respectively (Fig. 2C).

Figure 2. 

Descriptions of Pristimantis species across countries A number of species descriptions per country B number of annual species descriptions by country C number of annual species descriptions in the four countries with the highest species richness.

In total, 320 researchers are formally recognized as authors and co-authors in publications describing new species of Pristimantis (Suppl. material 1). Almost half (46%) of them have described a single species, while another 46% have described between two and seven species, 6.5% of authors have described 8–30 species, and finally only 0.6% of authors have described more than 50 species (Fig. 3A). Lynch is the author with the highest count of newly described species, with 195 species descriptions to his name. He is followed by Duellman with 82, and E. Lehr and M. Yánez-Muñoz with 36 each. Details on the 20 authors with the highest count of newly described species are provided in Table 2. Across all scientific collections where the type material has been deposited, the four institutions with the highest number of holotypes are the Museum of Natural History of University of Kansas (KU), Museo de Historia Natural del Instituto de Ciencias Naturales de Universidad Nacional de Colombia (ICN), Museo de Zoología de la Pontificia Universidad Católica del Ecuador (QCAZ), and División de Herpetología, Instituto Nacional de Biodiversidad, former Museo Ecuatoriano de Ciencias Naturales (DHMECN). KU and ICN house 35% of the typical material, and QCAZ and DHMECN have the 12.5% of type specimens of the genus (Fig. 3B). The rest of the collections hold a lower proportion of specimens than those mentioned (Fig. 3B, Suppl. material 1).

Figure 3. 

A Number of species described by the number of authors B number of holotypes of Pristimantis housed in the main scientific collections of natural history (an expanded list can be found in Suppl. material 1).

We identified a variety of eight languages used for the description of new Pristimantis species (Fig. 4A, Table 3), with the most common ones being English and Spanish. The English language has dominated species descriptions since the 1960s. The number of descriptions published in Spanish started to increase during the 1980s, though it remains lower than those published in English (Fig. 4A, Table 3). Conversely, there are more than 20 nationalities of researchers who have participated as authors and co-authors in descriptions of new species of Pristimantis (Fig. 4B). Researchers from the USA have significantly dominated, not only the total number of descriptions, but also the number of annual descriptions until 2002. As of this year, the participation of Ecuadorian authors has increased in greater numbers compared to their Colombian, Peruvian, and Venezuelan peers (Fig. 4B). The greatest percentage of authors involved in descriptions of new Pristimantis species are of Ecuadorian nationality (20.9%), followed by US Americans (18.8%) and Colombians (15.3%) (Table 4).

Figure 4. 

Languages and nationalities of authors who have participated in the descriptions of Pristimantis species A main languages used in descriptions per year B main nationalities of authors involved in descriptions per year.

From a gender perspective, 80% of the authors who have participated in the descriptions are male researchers and 20% female researchers (Figs 5, 6). In spite of an increase in the number of descriptions involving women starting in the 1950s, the participation of women in the description process continues to be considerably lower than that of their male counterparts (Fig. 5A). Ecuador is the country with the highest percentage of descriptions that incorporate female researchers with almost 18.8%, while Colombia, Peru, Venezuela, and Brazil have lower percentages (Fig. 5B). The bias towards male authors is overwhelmingly disproportionate in terms of principal authorship (i.e., first author or corresponding author). Where 50% of male authors featured as main authors, only 2% of female authors have held this role (Fig. 6, Table 5). We detected a significant slope between the proportion of female authors who participated in descriptions of all Pristimantis and the years (estimate = 0.03, SE = 0.01, t = 2.2, p = 0.02; Fig. 7). Of the four countries with the highest number of Pristimantis species descriptions, Ecuador was the only one with a significant positive slope between the proportion of female authors and time (estimate = 0.09, SE = 0.02, t = 3.8, p = < 0.001), while that on the contrary Colombia (estimate = 3.7 *10-3, SE = 0.01, t = 0.2, p = 0.8), Peru (estimate = -0.01, SE = 0.02, t = 3.8, p = 0.5),Venezuela (estimate = -0.01, SE = 0.03, t = -0.5, p = 0.7), and Brazil (estimate = -0.04, SE = 0.02, t = -2.25, p = 0.06) did not show any correlation between these variables (Fig. 7).

Figure 5. 

Male and female contributions to Pristimantis descriptions A number of female and male authors that participated in descriptions per year B contributions of female and male authors to descriptions per country.

Figure 6. 

Contributions in which authors have been principal author and defined as either first or corresponding author.

Figure 7. 

Change in the proportion of female authors over time. Gray shadow represents the confidence intervals of the Generalized Linear Model of the total proportion of female researchers (red solid line). Solid lines depict significant GLMs.

In relation to peer-reviewed journals in which the descriptions of new species of the genus have been published, Zootaxa and the Revista de la Academia Colombiana de Ciencias, Exactas, Físicas y Naturales have published the highest number of descriptions with 20.6% of the total described species. Other journals such as Herpetologica, ZooKeys, and Miscellaneous Publication of Museum of Natural History of University of Kansas have published 15.7% of the descriptions of Pristimantis (Fig. 8A). From the beginning of the descriptions of this diverse genus until the first decade of the 2000s, descriptions have been based mainly on morphology. Phylogenetic analyses were gradually incorporated into descriptions over the last 12 years, leading to a significant reduction in morphology-only descriptions of new Pristimantis species (Fig. 8B).

Figure 8. 

A Main scientific journals where descriptions of new Pristimantis species are published B type of descriptions per year, M = Morphological description, MP = morphological and phylogenetic description.

Regarding the conservation status of Pristimantis species, 24% are categorized by the IUCN Reed List as Least Concern, 31% are threatened (i.e., CR, EN, or VU), and 36.8% are Not Evaluated or Data Deficient (Fig. 9A). The country with the highest number of Not evaluated species is Ecuador (i.e., 39% of all Ecuadorian species). Peru and Venezuela host the highest percentages of species under the Data Deficient category, 31.8% and 42.8% of their total species, respectively. In Ecuador and Colombia, at least 30% of Pristimantis species are under some form of threat (Fig. 9B).

Figure 9. 

Conservation status categories established by the IUCN Red List for Pristimantis species A number of species per category B number of species in each category per country.

We are extremely grateful to Gorki Ríos-Alvear, David Brito-Zapata, and Maria Antonia Izurieta for their help in entering information in the database of Pristimantis descriptions. This research was supported by Universidad San Francisco de Quito USFQ through COCIBA Grants (project HUBI ID 12268, 17475) to CRP. We greatly appreciate the comments of Edgar Lehr and Juan Manuel Guayasamin, which helped improve the manuscript.