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| (A) Craugastor aenigmaticus (UCR 22961) from Cerro Arbolado, Puntarenas, CR, (B) C. blairi (SMF 104032) from Fortuna, PA, (C) C. sagui (SMF 104018) from La Nevera, PA, (D) C. zunigai (UCR 20389) from Potrero Grande, Puntarenas, CR, (E-F) C. podiciferus (UCR 23155, 23159) from Caribbean slopes of Cerro Kamuk, Limón, CR, (J) Craugastor sp. Pico Blanco (UCR 24466) from Escazú, San José, CR, (K) Craugastor sp. Chumacera (UCR 23011) from Chumacera, San José, CR, (L) Craugastor sp. Siola (UCR 23169) from Siola, Limón, CR, (M) C. bransfordii from Siquirres, Limón, CR, (O) C. underwoodi from Cascajal, San José, CR, (V) C. rearki from Siquirres, Limón, CR. in Arias, Crawford, Hertz et Olea,. 2025. |
Abstract
The Craugastor podiciferus Species Group contains eleven species of terraranan frogs distributed from eastern Honduras to eastern Panama. All species have remarkable color pattern polymorphisms, which may contribute to potential taxonomic problems. We performed exhaustive sampling throughout the geographic distribution of the group to evaluate the phylogenetic relationships and biogeographic history of all named species based on two mitochondrial markers and nuclear ddRAD loci. We also implemented various species delimitation methods to test for the presence of unconfirmed candidate species within the group. Molecular phylogenetic analyses showed that the group contains four major clades. All currently named species are supported by molecular data, yet species richness within the group is clearly underestimated. Species delimitation was discordant between the mitochondrial and nuclear datasets and among analytical methods. Adopting a conservative approach, we propose that the C. podiciferus species group contains at least 12 unconfirmed candidate species. Ancestral area reconstruction showed that the group originated and diversified in the highlands of the Talamancan montane forest ecoregion of Costa Rica and western Panama.
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| Photographs in life of Craugastor podiciferus species complex. (A) Craugastor aenigmaticus (UCR 22961) from Cerro Arbolado, Puntarenas, CR, (B) C. blairi (SMF 104032) from Fortuna, PA, (C) C. sagui (SMF 104018) from La Nevera, PA, (D) C. zunigai (UCR 20389) from Potrero Grande, Puntarenas, CR, (E-F) C. podiciferus (UCR 23155, 23159) from Caribbean slopes of Cerro Kamuk, Limón, CR, (G) Craugastor sp. Monte Verde (UCR 24613) from Monte Verde, Puntarenas, CR, (H) Craugastor sp. San Gerardo (CRARC 0247) from San Gerardo, Guanacaste, CR, (I) Craugastor sp. Fila Costeña (UCR 23028) from Quebradas, San José, CR, (J) Craugastor sp. Pico Blanco (UCR 24466) from Escazú, San José, CR, (K) Craugastor sp. Chumacera (UCR 23011) from Chumacera, San José, CR, (L) Craugastor sp. Siola (UCR 23169) from Siola, Limón, CR, (M) C. bransfordii from Siquirres, Limón, CR, (N) Craugastor sp. Fila Carbon (UCR 23127) from Amubri, Limón, CR, (O) C. underwoodi from Cascajal, San José, CR, (P) Craugastor sp. Quebradas from Fila Costeña, Puntarenas, CR, (Q) Craugastor sp. Vereh (UCR 23040) from Vereh, Cartago, CR, (R) Craugastor sp. Panama (SMF 104010) from Rambala. PA, (S) C. stejnegerianus (UCR 22976) from Palmar Norte, Puntarenas, CR, (T) C. gabbi (UCR 22998) from San Vito, Puntarenas, CR, (U) C. persimilis from Siquirres, Limón, CR, (V) C. rearki from Siquirres, Limón, CR, (W) Craugastor sp. Neilly (UCR 22985) from Río Claro, Puntarenas, CR, and (X) Craugastor sp. Quepos (UCR 24612) from Montes de Oca, San José CR. Photos by E. Arias (A, E, F, G, I, J, K, L, N, Q, S, T, W, and X), Andreas Hertz (B,C, and R), Eduardo Boza-Oviedo (D and O), Brian Kubicki (H, M, U, and V), and Raby Nuñez (P). |
Conclusions:
The diversity within the Craugastor podiciferus Species Group is vastly underestimated, as revealed by the presence of several undescribed species recovered from the phylogenetic and species delimitation analyses. An exhaustive morphological review of the genetic lineages may show morphological characteristics that would allow for the differentiation of the molecular lineages. Comprehensive studies are needed on habitat use, acoustics, behavior, and other data to corroborate and better understand the taxonomy of all lineages revealed here.
Based on our mitochondrial and nuclear analyses, morphological evidence, and previous information we recovered 23 lineages, 11 with names and 12 unconfirmed candidate species. Based on our results, we propose the following changes:
• We restrict C. podiciferus to populations of Cordillera Volcánica Central from Costa Rica and the Cordillera de Talamanca of Costa Rica and western Panama. In the Cordillera de Talamanca, C. podiciferus is restricted to the Caribbean slopes.
•Craugastor polyptychus is referred to as a junior synonym of C. bransfordii.
• Craugastor rearki is resurrected to include wide-ranging populations from the Caribbean versant of Costa Rica, Nicaragua, and Honduras.
• Craugastor lauraster is referred to as a junior synonym of the older name, C. rearki.
Finally, we want to highlight the need to continue exploring remote areas in the ICA, especially in the Talamanca Mountain Range. The fieldwork performed in this area has resulted in the discovery of several new species or new records for the region. Therefore, more fieldwork and laboratory work are necessary to improve the knowledge of biodiversity in this region to perform informed strategies of conservation.
Erick Arias, Andrew J. Crawford, Andreas Hertz and Gabriela Parra Olea. 2025. Deep Cryptic Diversity in the Craugastor podiciferus Species Group (Anura: Craugastoridae) of Isthmian Central America revealed by mitochondrial and nuclear data. PeerJ. 13:e18212. DOI: doi.org/10.7717/peerj.18212
