-
摘要: 本文结合分子、形态和鸣声数据,描述巴拿马东部斑蟾属Atelopus一新种,即A. fronterizo sp. nov. 。新种可依据以下形态特征与同属近缘物种进行区分:1)第一指微弱;2)雌性SVL 35.1–50.1mm(n=13),HW/SVL 0.23–0.34 (n=59), EYND/HW 0.27–0.39 (n=60), TIBL/SVL 0.41–0.56 (n=58), 以及 HAL/SVL 0.22–0.28 (n=49);3)背部底色黄色或绿色,伴有密集的深橄榄色横纹或斑块;4)求偶鸣声长176–235ms,伴有19–34个脉冲,平均脉冲重复率为131.69脉冲/秒,鸣声主频率2422.50–2606.50Hz。新种嵌于中美洲的斑蟾支系中,其与近缘种A. certus和A. glyphus在16S和COI线粒体片段上的最小K2P遗传距离均大于2.6%和4.9%。依据16S和COI片段构建的贝叶斯和极大似然法系统演化关系一致,节点支持率较高。依据上述线粒体片段重建的单倍型网络也支持新种自成一支,种内突变步数较低(16S单倍型1-4步,COI单倍型1步),而相比近缘种而言,最低突变步数则普遍较高,其中16S单倍型较A. certus为9步,COI单倍型较A. glyphus为28步。Abstract: A new species of the genus Atelopus, Atelopus fronterizo sp. nov. , from eastern Panama is described herein based on molecular, morphological, and bioacoustic evidence. The new species can be distinguished from its congeners occurring in the region by a combination of the following characters: (1) phalangeal reduction in thumb; (2) SVL (females only) (35.1–50.1; n=13), HW/SVL (0.23–0.34; n=59), EYND/HW (0.27–0.39; n=60), TIBL/SVL (0.41–0.56; n=58), and HAL/SVL (0.22–0.28; n=49); (3) dorsal color pattern with green or yellow background and extensive dark olive blotches forming transversal bands or mottling; (4) advertisement call duration 176–235 ms with 19–34 pulses, average pulse rate 131.69 pulses/s, and dominant frequency 2 422.50–2 606.50 Hz. The new species is nested within the Central American clade of Atelopus. The minimum Kimura‐2‐parameter (K2P) genetic divergence between Atelopus fronterizo sp. nov. and its most phylogenetically similar congeners (A. certus and A. glyphus) is >2.6% for 16S and >4.9% for COI (
Table 1 ). The phylogenetic relationship is strongly supported by ultrafast bootstrap values for the maximum-likelihood trees of both genetic markers (16S, 96; COI, 100,Figure 1A ). Bayesian analysis of the concatenated sequences resulted in a tree with similar topology and high posterior probability support (0.99; Supplementary Figure S1). In addition, haplotype networks inferred from COI and 16S (Supplementary Figure S2) showed a well-separated clade containing the new species (two for COI, four for 16S). The number of mutational steps between haplotypes for the new species samples is very low (1–4 in 16S; one in COI), and the minimum number of mutational steps from the nearest species is nine for 16S (distance to A. certus) and 28 for COI (distance to A. glyphus).-
Key words:
- Integrative taxonomy /
- Darién /
- Endangered species /
- Harlequin frog /
- New species
-
Figure 1. Molecular comparison and geographic distribution of Panamanian species of Atelopus, and main features of holotype of Atelopus fronterizo sp. nov.
A: Maximum-likelihood trees based on COI and 16S sequences of Atelopus species from Panamá. Colors of branches indicate bootstrap support, values >90 are marked on branches. B: Map of Panama showing distribution of Atelopus species occurring in the country; C: Atelopus fronterizo sp. nov. holotype (MHCH 3110), dorsal view of preserved specimen; D: Ventral view; E: Head laterally; F: Left hand; G: Left foot; H: Live holotype, from Púcuro River (MHCH 3110); scale bars correspond to 5 mm. All photos were taken by Abel Batista, except for that of A. zeteki, taken by Emanuele Biggi.
Table 1. K2P distances for mitochondrial fragments of 16S and COI genes of Atelopus species from Panamá and Colombian samples of A spurrelli.
Species 1 2 3 4 5 6 7 n=2 n=2 n=3 n=1 n=7 n=19 1 A. fronterizo sp. nov. 0.24 5.18–5.69 4.93–5.70 8.88–9.16 8.64–10.02 9.74–11.49 n=4 0.50–2.04 2 A. certus 0.71 2.41–3.14 9.48–9.74 9.98 - 10.02 9.98–10.57 n=2 2.55–3.08 1.01 3 A. glyphus 0.00–0.47 8.88–9.14 9.18–9.74 8.64–9.72 n=3 2.55–3.60 1.52–2.55 0.00–0.50 4 A. spurrelli 0.00 8.04–8.85 8.85–11.71 n=2 4.67–5.22 5.75 5.74–6.29 0.00 5 A. varius n=5 3.61–4.67 4.67–5.75 4.15–5.22 4.12–4.65 0.00–0.50 6 A. zeteki 0.00–3.90 0.00–6.50 n=7 3.60–5.77 4.66–6.87 4.13–6.33 4.11–5.74 0.00–2.03 0.00–2.55 7 A. limosus 0.00–6.23 n=23 4.15–5.22 4.67–6.31 4.15–5.77 4.12–5.19 0.50–2.03 0.50–3.08 0.00–1.52 8 A. chiriquiensis n=1 4.15–4.69 4.13–4.67 3.61–4.15 4.12 1.52–2.03 1.52–3.08 1.52–2.04 Ranges represent minimum and maximum observed distances. Intraspecific genetic distances are shown along diagonal of matrix. Values correspond to interspecific genetic distances of 16S and COI genes (COI in bold); COI sequences of A. varius and A. chiriquiensis were not available. -
[1] Acosta-Galvis AR, Saldarriaga-Gómez AM, Ramírez B, Vargas-Ramírez M. 2020. A new Terrarana frog of genus Pristimantis from an unexplored cloud forest from the eastern Andes, Colombia. ZooKeys, 961: 129−156. doi: 10.3897/zookeys.961.51971 [2] AmphibiaWeb. 2020[2020-10-14]. Berkeley, CA, USA: University of California, https://amphibiaweb.org. [3] Barbour T. 1923. Notes on reptiles and amphibians from Panama. Occasional Papers, 129. Michigan: University of Michigan, 1–16. [4] Batista A, Hertz A, Köhler G, Mebert K, Veselý M. 2014a. Morphological variation and phylogeography of frogs related to Pristimantis caryophyllaceus (Anura: Terrarana: Craugastoridae) in Panama. Salamandra, 50(3): 155−171. [5] Batista A, Hertz A, Mebert K, Köhler G, Lotzkat S, Ponce M, et al. 2014b. Two new fringe-limbed frogs of the genus Ecnomiohyla (Anura: Hylidae) from Panama. Zootaxa, 3826(3): 449−474. doi: 10.11646/zootaxa.3826.3.2 [6] Batista A, Köhler G, Mebert K, Hertz A, Veselý M. 2016. An integrative approach to reveal speciation and species richness in the genus Diasporus (Amphibia: Anura: Eleutherodactylidae) in eastern Panama. Zoological Journal of the Linnean Society, 178(2): 267−311. doi: 10.1111/zoj.12411 [7] Batista A, Köhler G, Mebert K, Veselý M. 2014c. A new species of Bolitoglossa (Amphibia: Plethodontidae) from eastern Panama, with comments on other species of the adspersa species group from eastern Panama. Mesoamerican Herpetology, 1(1): 97−121. [8] Bravo-Valencia L, Rivera-Correa M. 2011. A new species of harlequin frog (Bufonidae: Atelopus) with an unusual behavior from Andes of Colombia. Zootaxa, 3045(1): 57−67. doi: 10.11646/zootaxa.3045.1.4 [9] Breder CM. 1946. Amphibians and reptiles of the Rio Chucunaque drainage, Darién, Panamá, with notes on their life histories and habits. Bulletin of the American Museum of Natural History, 86(8): 375−436. [10] Chambers EA, Hebert PD. 2016. Assessing DNA barcodes for species identification in North American reptiles and amphibians in natural history collections. PLoS One, 11(4): e0154363. doi: 10.1371/journal.pone.0154363 [11] Cocroft RB, McDiarmid RW, Jaslow AP, Ruiz-Carranza PM. 1990. Vocalizations of eight species of Atelopus (Anura: Bufonidae) with comments on communication in the genus. Copeia, 1990(3): 631−643. doi: 10.2307/1446428 [12] Coloma LA, Lötters S, Salas AW. 2000. Taxonomy of the Atelopus ignescens complex (Anura: Bufonidae): designation of a neotype of Atelopus ignescens and recognition of Atelopus exiguus. Herpetologica, 56(3): 303−324. [13] Crawford AJ, Lips KR, Bermingham E. 2010. Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama. Proceedings of the National Academy of Sciences of the United States of America, 107(31): 13777−13782. doi: 10.1073/pnas.0914115107 [14] De la Riva I, Castroviejo-Fisher S, Chaparro JC, Boistel R, Padial JM. 2011. A new species of Atelopus (Anura: Bufonidae) from the Amazonian slopes of the Andes in south-eastern Peru. Salamandra, 47(3): 161−168. [15] Dunn ER. 1931. New frogs from Panama and Costa Rica. Occasional papers of the Boston Society of Natural History, 5: 385−401. [16] Dunn ER. 1933. Amphibians and reptiles from El Valle de Antón, Panamá. Occasional Papers of the Boston Society of Natural History, 8: 65−79. [17] Flechas SV, Blasco-Zúñiga A, Merino-Viteri A, Ramírez-Castañeda V, Rivera M, Amézquita A. 2017a. The effect of captivity on the skin microbial symbionts in three Atelopus species from the lowlands of Colombia and Ecuador. PeerJ, 5: e3594. doi: 10.7717/peerj.3594 [18] Flechas SV, Paz A, Crawford AJ, Sarmiento C, Acevedo AA, Arboleda A, et al. 2017b. Current and predicted distribution of the pathogenic fungus Batrachochytrium dendrobatidis in Colombia, a hotspot of amphibian biodiversity. Biotropica, 49(5): 685−694. doi: 10.1111/btp.12457 [19] Frost D. 2020[2020-09-20]. Amphibian species of the world 6.1, an online reference. https://amphibiansoftheworld.amnh.org/index.php. [20] Guayasamin JM, Bonaccorso E, Duellman WE, Coloma LA. 2010. Genetic differentiation in the nearly extinct harlequin frogs (Bufonidae: Atelopus), with emphasis on the Andean Atelopus ignescens and A. bomolochos species complexes. Zootaxa, 2574(1): 55−68. doi: 10.11646/zootaxa.2574.1.2 [21] Heckadon-Moreno S. 1996. Estudios Botánicos y etnográficos de Henri Pittier en la costa de San Blas y Puerto Obaldía – 1911. Épocas: Segunda Era, 11(10): 10−11. [22] Hogan CM, World Wildlife Fund. 2014[2020-05-25]. Chocó-Darién moist forests. http://editors.eol.org/eoearth/wiki/Chocó-Darién_moist_forests. [23] Ibáñez DR, Jaramillo CA, Solís FA. 1995. Una especie nueva de Atelopus (Amphibia: Bufonidae) de Panama. Caribbean Journal of Science, 31(1–2): 57−64. [24] Jaslow AP. 1979. Vocalization and aggression in Atelopus chiriquiensis (Amphibia, Anura, Bufonidae). Journal of Herpetology, 13(2): 141−145. doi: 10.2307/1563919 [25] Jorge RF, Ferrão M, Lima AP. 2020. Out of Bound: A new threatened harlequin toad (Bufonidae, Atelopus) from the outer borders of the Guiana shield in Central Amazonia described through integrative taxonomy. Diversity, 12(8): 310. doi: 10.3390/d12080310 [26] Kimura M. 1980. A simple method for estimating evolutionary rates of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16(2): 111−120. doi: 10.1007/BF01731581 [27] La Marca E, Lips KR, Lötters S, Puschendorf R, Ibáñez R, Rueda-Almonacid JV, et al. 2005. Catastrophic population declines and extinctions in Neotropical harlequin frogs (Bufonidae: Atelopus). Biotropica, 37(2): 190−201. doi: 10.1111/j.1744-7429.2005.00026.x [28] Lewis CHR, Richards-Zawacki CL, Ibáñez R, Luedtke J, Voyles J, Houser P, et al. 2019. Conserving Panamanian harlequin frogs by integrating captive-breeding and research programs. Biological Conservation, 236: 180−187. doi: 10.1016/j.biocon.2019.05.029 [29] Lichtenstein H, von Martens E. 1856. Nomenclator Reptilium et Amphibiorum Musei Zoologici Berolinensis. Namenverzeichniss der in der zoologischen Sammlung der Königlichen Universität zu Berlin aufgestellten Arten von Reptilien und Amphibien nach ihren Ordnungen, Familien und Gattungen. Berlin: Königliche Akademie der Wissenschaften. [30] Lips KR, Brem F, Brenes R, Reeve JD, Alford RA, Voyles J, et al. 2006. Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community. Proceedings of the National Academy of Sciences of the United States of America, 103(9): 3165−3170. doi: 10.1073/pnas.0506889103 [31] Lötters S. 1996. The Neotropical Toad Genus Atelopus. Checklist—Biology—Distribution. Cologne: Vences and Glaw Verlags GbR. [32] Lötters S, Mebs D, Köhler G, Vargas J, La Marca E. 2019. The voice from the hereafter: vocalisations in three species of Atelopus from the Venezuelan Andes, likely to be extinct. Herpetozoa, 32: 267−275. doi: 10.3897/herpetozoa.32.e39192 [33] Lötters S, van der Meijden A, Coloma LA, Boistel R, Cloetens P, Ernst R, et al. 2011. Assessing the molecular phylogeny of a near extinct group of vertebrates: the Neotropical harlequin frogs (Bufonidae; Atelopus). Systematics and Biodiversity, 9(1): 45−57. doi: 10.1080/14772000.2011.557403 [34] Myers CW, Duellman WE. 1982. A new species of Hyla from Cerro Colorado, and other tree frog records and geographical notes from western Panama. American Museum Novitates, 2752: 1−32. [35] Nagy ZT, Sonet G, Glaw F, Vences M. 2012. First large‐scale DNA barcoding assessment of reptiles in the biodiversity hotspot of Madagascar, based on newly designed COI primers. PLoS One, 7(3): e34506. doi: 10.1371/journal.pone.0034506 [36] OSM Contributors. 2015[2019-02-25]. https://planet.openstreetmap.org [37] Ramírez JR, Jaramillo CA, Lindquist ED, Crawford AJ, Ibáñez R. 2020. Recent and rapid radiation of the highly endangered Harlequin frogs (Atelopus) into Central America inferred from mitochondrial DNA sequences. Diversity, 12(9): 360. doi: 10.3390/d12090360 [38] Rivera-Correa M. 2005. Atelopus spurrelli. In: Rueda-Almonacid JV, Rodríguez-Mahecha JV, La Marca E, Lötters S, Kahn T, Angulo A. Ranas Arlequines. Bogotá. Conservación Internacional, 158. [39] Sabaj MH. 2016(2016-08-16). Standard symbolic codes for institutional resource collections in herpetology and ichthyology: an online reference. Version 6.5. Washington, DC: American Society of Ichthyologists and Herpetologists. http://www.asih.org/. [40] Savage JM. 1972. The harlequin frogs, genus Atelopus, of Costa Rica and western Panama. Herpetologica, 28(2): 77−94. [41] Savage JM. 2002. The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. Chicago: The University of Chicago Press. [42] Savage JM, Heyer WR. 1997. Digital webbing formulae for anurans: a refinement. Herpetological Review, 28(3): 131. [43] Shreve B. 1936. A new Atelopus from Panama and a new Hemidactylus from Colombia. Occasional Papers of the Boston Society of Natural History, 8: 269−272. [44] Vieites DR, Wollenberg KC, Andreone F, Köhler J, Glaw F, Vences M. 2009. Vast underestimation of Madagascar’s biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the United States of America, 106(20): 8267−8272. doi: 10.1073/pnas.0810821106 [45] World Wildlife Fund. 2014(2014-05-14)[2020-05-25]. Eastern Panamanian montane forests. http://editors.eol.org/eoearth/wiki/Eastern_Panamanian_montane_forests. [46] Zippel KC, Ibáñez R, Lindquist ED, Richards CL, Jaramillo CA, Griffith EJ. 2006. Implicaciones en la conservación de las ranas doradas de Panamá, asociadas con su revisión taxonómica. Herpetotropicos, 3(1): 29−39. -
ZR-2020-319 Supplementary materials.pdf
-