Skip to main content Skip to main navigation menu Skip to site footer
Type: Monograph
Published: 2025-01-21
Page range: 1-76
Abstract views: 357
PDF downloaded: 12

A roadmap for harlequin frog systematics, with a partial revision of Amazonian species related to Atelopus spumarius

Biogeography Department; Trier University; 54286 Trier; Germany
Biogeography Department; Trier University; 54286 Trier; Germany
Département Homme & Environnement; Muséum national d’Histoire naturelle; CNRS; 75005 Paris; France
Fundación Atelopus; Santa Marta; Colombia
Laboratory Mecadev UMR 7179 CNRS/MNHN; Département Adaptations du Vivant; Bâtiment d’Anatomie Comparée; 75005 Paris; France
Department of Biological Sciences; Florida International University; Miami; FL 33199; USA
Museo de Biodiversidad del Perú; Cusco; Peru
Instituto Peruano de Herpetología (IPH); Lima; Peru
Dendrobirds Peru; Tarapoto; Peru
Physiologie moléculaire et adaptation; Muséum national d’Histoire naturelle; CNRS; 75231 Paris; France
Centro Jambatu de Investigación y Conservación de Anfibios; Fundación Jambatu; San Rafael; Ecuador
Department of Biological Sciences; Universidad de los Andes; Bogotá; Colombia
Photo Wildlife Tours; Quito; Ecuador; Fundación Cóndor Andino; Quito; Ecuador
Museo de Historia Natural; Universidad Nacional Mayor de San Marcos; Lima; Peru
Instituto de Biología; Universidad de Antioquia; Medellín; Colombia
Museo Nacional de Ciencias Naturales-CSIC; Madrid; Spain
Biogeography Department; Trier University; 54286 Trier; Germany
Senckenberg Natural History Collections Dresden; Museum of Zoology; 01109 Dresden; Germany; Technical University of Dresden; Faculty of Biology; Dresden; Germany
Bichos.team; Bogotá; Colombia
Centre de Recherche su la Biodiversité et l´Environnment (CRBE) - UMRS 5300; Université Toulouse III Paul Sabatier; 31062 Toulouse cedex 9; France
Universidad San Francisco de Quito USFQ; Colegio de Ciencias Biológicas y Ambientales COCIBA; Laboratorio de Biología Evolutiva; Quito; Ecuador; Tandayapa Cloud Forest Station; Universidad San Francisco de Quito USFQ; Quito; Ecuador
Biogeography Department; Trier University; 54286 Trier; Germany
Laboratório de Ecologia e Conservação de Populações (LECP); Programa de Pós-graduação em Ecologia; Universidade Federal do Rio de Janeiro (UFRJ); Rio de Janeiro; RJ; Brazil
Biogeography Department; Trier University; 54286 Trier; Germany
Senckenberg Natural History Collections Dresden; Museum of Zoology; 01109 Dresden; Germany
Senckenberg Natural History Collections Dresden; Museum of Zoology; 01109 Dresden; Germany
Biogeography Department; Trier University; 54286 Trier; Germany
Escuela de Geografía; Facultad de Ciencias Forestales y Ambientales; Universidad de Los Andes; Mérida 5101; Venezuela
Fundación para el Desarrollo de las Ciencias Físicas; Matemáticas y Naturales; Caracas; Venezuela; Instituto Venezolano de Investigaciones Científicas (FUDECI); Caracas; Venezuela
Grupo de Morfología y Ecología Evolutiva; Instituto de Ciencias Naturales; Universidad Nacional de Colombia; Bogotá D.C.; Colombia
Viks byvag 17; Norrtalje; Sweden
Department of Biology; University of Dayton; Dayton; Ohio; USA
Fundación Atelopus; Santa Marta; Colombia
Synchrotron SOLEIL L’Orme des Merisiers; Départementale 128; 91190 Saint-Aubin; France
Centro Jambatu de Investigación y Conservación de Anfibios; Fundación Jambatu; San Rafael; Ecuador
Fundación Ecominga; Quito; Ecuador
Instituto Peruano de Herpetología (IPH); Lima; Peru
Zukunftskolleg; University of Konstanz; Konstanz; Germany; Department of Ecology of Animal Societies; Max Planck Institute of Animal Behavior; Konstanz; Germany
Fundación Atelopus; Santa Marta; Colombia; Grupo de Investigación en Biodiversidad y Ecología Aplicada (GIBEA); Facultad de Ciencias Básicas; Universidad del Magdalena; Santa Marta; Magdalena; Colombia
Centro de Investigación de la Biodiversidad y Cambio Climático (BioCamb) e Ingeniería en Biodiversidad y Recursos Genéticos; Facultad de Ciencias de Medio Ambiente; Universidad Indoamérica; Av. Machala y Sabanilla; Quito; Ecuador
Estación Biológica de Doñana (EBD-CSIC); Avenida Américo Vespucio; 26; 41092 Seville; Spain
Laboratory Mecadev UMR 7179 CNRS/MNHN; Département Adaptations du Vivant; Bâtiment d’Anatomie Comparée; 75005 Paris; France
Centro Jambatu de Investigación y Conservación de Anfibios; Fundación Jambatu; San Rafael; Ecuador
Department of Biological Sciences; Universidad de los Andes; Bogotá; Colombia
Biogeography Department; Trier University; 54286 Trier; Germany
Instituto Peruano de Herpetología (IPH); Lima; Peru; Rainforest Partnership; 4005 Guadalupe St.; Austin; Texas 78751; USA
Biology Program; California State University Channel Islands; One University Drive; Camarillo; CA 93012; USA
Department of Biology; University of Dayton; Dayton; Ohio; USA
Biogeography Department; Trier University; 54286 Trier; Germany
Amphibia Amphibian crisis

Abstract

Harlequin frogs, genus Atelopus, are a species-rich group of bufonid anurans from the Neotropics with more than 100 species. For nearly four decades now, this group has suffered from massive population declines. Almost all species are threatened with extinction, and many populations and several species are considered extinct or possibly extinct. This results in a limited sampling available for studies on harlequin frog systematics, especially in terms of molecular genetic information. However, efficient conservation of harlequin frogs requires an improved taxonomy. This is further complicated through the circumstance that many Atelopus species are relatively poor in external morphological characters combined with a high level of intra-specific character variation (e.g. coloration and body size). At the same time, cryptic diversity exists with well differentiated species (supported by osteology and molecular genetics) almost indistinguishable by external morphology. We compiled the largest dataset to date for mitochondrial (12S, 16S, cyt b) and nuclear (POMC, RAG1) markers and present a phylogeny (likelihood and Bayesian inference methods) including 152 samples from 104 populations scattered over the entire geographic range of the genus. Four allo- or parapatric main clades are distinguished: I. Sierra Nevada; II. Venezuelan-Andean; III. Andean-Chocó-Central American (with the ignescens and the varius-longirostris clades); and IV Amazonian (containing the tricolor and the flavescens-spumarius clades). The phylogenetic relationships within these clades remain to be resolved. Taxonomic implications included both splitting and lumping, but taxonomic action is here only taken for populations related to A. spumarius from western Amazonia. Besides redescriptions of A. spumarius sensu stricto and A. colomai, we describe two new species based on morphology, skull osteology and bioacoustics. Additional yet understudied populations from Amazonia may be allocated to these species or may represent additional undescribed taxa.

 

References

  1. Asquith, A. & Altig, R. (1987) Atelopus spumarius. Vocalization. Herpetological Review, 18, 32–33.
  2. Ballestas, O., Lampo, M. & Rodríguez, D. (2021) Living with the pathogenic chytrid fungus: Exploring mechanisms of coexistence in the harlequin toad Atelopus cruciger. PLoS ONE, 16, e0254439. https://doi.org/10.1371/journal.pone.0254439
  3. Barbour, T. (1905) The vertebrata of Gorgona Island, Colombia: Reptilia; Amphibia. Bulletin of the Museum of Comparative Zoology, 46, 98–102.
  4. Benson, D.A., Clark, K., Karsch-Mizrachi, I., Lipman, D.J., Ostell, J. & Sayers, E.W. (2015) GenBank. Nucleic Acids Research, 43, D30. https://doi.org/10.1093/nar/gku1216
  5. Biton, R., Boistel, R., Rabinovich, S.G., Gafny, S., Brumfeld, V. & Bailon, S. (2016) Osteological observations on the Alytid Anura Latonia nigriventer with comments on functional morphology, biogeography, and evolutionary history. Journal of Morphology, 277, 1131–1145. https://doi.org/10.1002/jmor.20562
  6. Boistel, R., Grosjean, S. & Lötters, S. (2005) Tadpole of Atelopus franciscus from French Guyana, with comments on other larvae of the genus (Anura: Bufonidae). Journal of Herpetology, 39, 148–153. https://doi.org/10.1670/0022-1511(2005)039[0147:TOAFFF]2.0.CO;2
  7. Boistel, R., Swoger, J., Kržič, U., Fernandez, V., Gillet, B. & Reynaud, E.G. (2011) The future of three‐dimensional microscopic imaging in marine biology. Marine Ecology, 32, 438–452. https://doi.org/10.1111/j.1439-0485.2011.00442.x
  8. Boistel, R., Aubin, T., Cloetens, P., Peyrin, F., Scotti, T., Herzog, P., Gerlach, J., Pollet, N. & Aubry, J.-F. (2013) How minute sooglossid frogs hear without a middle ear. Proceedings of the Academy of Natural Sciences of Philadelphia, 110 (38), 15360–15364. https://doi.org/10.1073/pnas.1302218110
  9. Boulenger, G.A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. 2nd Edition. Taylor and Francis, London, 503 pp.
  10. Böning, P., Lötters, S., Barzaghi, B., Bock, M., Bok, B., Bonato, L., Ficetola, G.F., Glaser, F., Griese, J., Grabher, M., Leroux, C., Gopikrishna, M., Manenti, R., Ludwig, G., Preininger, D., Rödel, M.-O., Seibold, S., Smith, S., Tiemann, L., Thein, J., Veith, M. & Plewnia, A. (2024) Alpine salamanders at risk? The current status of an emerging fungal pathogen. PLoS ONE, 19, e0298591. https://doi.org/10.1371/journal.pone.0298591
  11. Bravo-Valencia, L. & Rivera-Correa, M. (2011) A new species of harlequin frog (Bufonidae: Atelopus) with an unusual behaviour from Andes of Colombia. Zootaxa, 3045 (1), 57–67. https://doi.org/10.11646/zootaxa.3045.1.4
  12. Bush, M.B. (1994) Amazonian speciation: a necessarily complex model. Journal of Biogeography, 21, 5–17. https://doi.org/10.2307/2845600
  13. Byrne, A.Q., Richards-Zawacki, C.L., Voyles, J., Bi, K., Ibáñez, R. & Rosenblum, E.B. (2020) Whole exome sequencing identifies the potential for genetic rescue in iconic and Critically Endangered Panamanian harlequin frogs. Global Change Biology, 27, 50–70. https://doi.org/10.1111/gcb.15405
  14. Cannatella, D.C. (1981) A new Atelopus from Ecuador and Colombia. Journal of Herpetology, 15, 133–138. https://doi.org/10.2307/1563371
  15. Carrillo-Bilbao, G. & Martin-Solano, S. (2013) Nuevo registro de distribución de Atelopus spumarius Cope, 1871 (Anura: Bufonidae) para Ecuador. Revista Latinoamericana de Conservación, 3, 48–50.
  16. Cocroft, R.B., McDiarmid, R.W., Jaslow, A.P. & Ruiz-Carranza, P.M. (1990) Vocalizations of eight species of Atelopus (Anura: Bufonidae) with comments on communication in the genus. Copeia, 1990, 631–643. https://doi.org/10.2307/1446428
  17. Coloma, L.A. (1997) Morphology, Systematics, and Phylogenetic Relationships among Frogs of the Genus Atelopus (Anura: Bufonidae). Doctoral Dissertation, University of Kansas, Lawrence, 267 pp.
  18. Coloma, L.A., Lötters, S. & Salas, A. (2000) Taxonomy of the Atelopus ignescens complex (Anura: Bufonidae): Designation of a neotype of Atelopus ignescens and recognition of Atelopus exiguus. Herpetologica, 56, 303–324.
  19. Coloma, L.A. (2002) Two new species of Atelopus (Anura: Bufonidae) from Ecuador. Herpetologica, 58, 229–252. https://doi.org/10.1655/0018-0831(2002)058[0229:TNSOAA]2.0.CO;2
  20. Coloma, L.A., Lötters, S., Duellman, W.E. & Miranda-Leiva, A. (2007) A taxonomic revision of Atelopus pachydermus, and description of two new (extinct?) species of Atelopus from Ecuador (Anura: Bufonidae). Zootaxa, 1557 (1), 1–32. https://doi.org/10.11646/zootaxa.1557.1.1
  21. Coloma, L.A., Duellman, W.E., Almendáriz C., A., Ron, S.R., Terán-Valdez, A. & Guayasamin, J.M. (2010) Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru. Zootaxa, 2574 (1), 1–54. https://doi.org/10.11646/zootaxa.2574.1.1
  22. Cope, E.D. (1871) Ninth contribution to the herpetology of Tropical America. Proceedings of the Academy of Natural Sciences of Philadelphia, 23, 200–224.
  23. Costa-Campos, C.E. & Carvalho, T. (2018) The advertisement call of the Hoogmoed’s harlequin toad Atelopus hoogmoedi Lescure, 1974 from northern Brazil (Anura, Bufonidae). Zootaxa, 4521 (1), 141–144. https://doi.org/10.11646/zootaxa.4521.1.11
  24. Crump, M.L. (1986) Homing and site fidelity in a Neotropical frog, Atelopus varius (Bufonidae). Copeia, 1986, 438–444.
  25. Culebras, J., Novales, A., Quezada Riera, A.B., Medina, D. & Plewnia, A. (2023) New records of the glassfrog Hyalinobatrachium iaspidiense (Ayarzagüena, 1992) in western Amazonia. Herpetology Notes, 16, 179–182. https://doi.org/10.2307/1445001
  26. Da Silva, G., Cornélio, G.S., de Oliveira, E.A., Trindade, N., França, I. & Hernández Ruz, E. (2020) A candidate species currently classified as Atelopus hoogmoedi (Anura: Bufonidae) in the eastern Amazon, Pará, Brazil. Genetics and Molecular Research, 19, gmr18392. https://doi.org/10.4238/gmr18392
  27. De la Riva, I., Castroviejo-Fisher, S., Chaparro, J.C., Boistel, R. & Padial, J.M. (2011) A new species of Atelopus (Anura: Bufonidae) from the Amazonian slopes of the Andes in south-eastern Peru. Salamandra, 47, 161–168.
  28. De Magalhães, R.F., Rocha, P.C., Santos, F.R., Strüssmann, C. & Giaretta, A.A. (2018) Integrative taxonomy helps to assess the extinction risk of anuran species. Journal for Nature Conservation, 45, 1–10. https://doi.org/10.1016/j.jnc.2018.07.001
  29. De Queiroz, K. (1998) The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In: Raman, S. (Ed.), Endless Forms: Species and Speciation. Oxford University Press, Oxford, pp. 57–75.
  30. Dole, J.W. & Durant, P. (1974) Movements and seasonal activity of Atelopus oxyrhynchus (Anura: Atelopodidae) in a Venezuelan Cloud Forest. Copeia, 1974, 230–235. https://doi.org/10.2307/1443028
  31. dos Santos Dias, P.H. & Anganoy-Criollo, M. (2024) Harlequin frog tadpoles—comparative buccopharyngeal morphology in the gastromyzophorous tadpoles of the genus Atelopus (Amphibia, Anura, Bufonidae), with discussion on the phylogenetic and evolutionary implication of characters. The Science of Nature, 111 (1), 3. https://doi.org/10.1007/s00114-024-01889-6
  32. Duellman, W.E. & Lynch, J.D. (1969) Descriptions of Atelopus tadpoles and their relevance to atelopodid classification. Herpetologica, 25, 231–240.
  33. Fabrezi, M. & Alberch, P. (1996) The carpal elements of anurans. Herpetologica, 52, 188–204.
  34. Flechas, S.V., Sarmiento, C. & Amézquita, A. (2012) Bd on the beach: high prevalence of Batrachochytrium dendrobatidis in the lowland forest of Gorgona Island (Colombia, South America). EcoHealth, 9, 298–302. https://doi.org/10.1007/s10393-012-0771-9
  35. Flechas, S.V., Vredenburg, V.T. & Amézquita, A. (2015) Infection prevalence in three lowland species of harlequin toads from the threatened genus Atelopus. Herpetological Review, 46, 528–532.
  36. Fouquet, A., Recoder, R., Teixeira Jr, M., Cassimiro, J., Amaro, R.C., Camacho, A., Damasceno, R., Carnaval, A.C., Moritz, C. & Rodrigues, M.T. (2012) Molecular phylogeny and morphometric analyses reveal deep divergence between Amazonia and Atlantic Forest species of Dendrophryniscus. Molecular Phylogenetics and Evolution, 62, 826–838. https://doi.org/0.1016/j.ympev.2011.11.023
  37. Frost, D.R. (2024) Amphibian species of the world: an online reference. Version 6.1. Available from: https://amphibiansoftheworld.amnh.org/ (accessed 21 April 2024)
  38. Frost, D.R. & Kluge, A.G. (1994) A consideration of epistemology in systematic biology, with special reference to species. Cladistics, 10, 259–294. [http://hdl.handle.net/2027.42/31356]
  39. Gascon, C. (1989) The tadpole of Atelopus pulcher Boulenger (Anura, Bufonidae) from Manaus, Amazonas. Revista Brasileira de Zoologia, 6, 235–239. https://doi.org/10.1590/S0101-81751989000200007
  40. Gosner, K.L. (1960) A simplified table for staging Anura embryos and larvae with notes on identification. Herpetologica, 16, 183–190.
  41. Gray, P. & Cannatella, D.C. (1985) A new species of Atelopus (Anura, Bufonidae) from the Andes of northern Perú. Copeia, 1985, 910–917. https://doi.org/10.2307/1445241
  42. Guayasamin, J.M., Bonaccorso, E., Duellman, W.E. & Coloma, L.A. (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. https://doi.org/10.11646/zootaxa.2574.1.2
  43. Guindon, S., Dufayard, J.-F., Lefort, V., Anisimova, M., Hordijk, W. & Gascuel, O. (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology, 59, 307–321. https://doi.org/10.1093/sysbio/syq010
  44. Hausdorf, B. (2011) Progress towards a general species concept. Evolution, 64, 923–931. https://doi.org/10.1111/j.1558-5646.2011.01231.x
  45. Herrera-Alva, V., Díaz, V., Castillo, E., Rodolfo, C. & Catenazzi, A. (2020) A new species of Atelopus (Anura: Bufonidae) from southern Peru. Zootaxa, 4853 (3), 404–420. https://doi.org/10.11646/zootaxa.4853.3.4
  46. Hoang, D.T., Chernomor, O., von Haeseler, A., Minh, B.Q. & Vinh L.S. (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution, 35, 518–522. https://doi.org/10.1093/molbev/msx281
  47. Ibáñez D., R., Jaramillo, C.A. & Solís, F.A. (1995) Una especie nueva de Atelopus (Amphibia: Bufonidae) de Panama. Caribbean Journal of Science, 31, 57–64.
  48. IUCN (2012) IUCN Red List Categories and Criteria. Version 3.1. 2nd Edition. IUCN, Gland, 32 pp.
  49. IUCN (2023) The IUCN Red List of Threatened Species. Version 2023-1. Available from: https://www.iucnredlist.org (accessed 22 December 2023)
  50. Jaynes, K.E., Páez-Vacas, M.I., Salazar-Valenzuela, D., Guayasamin, J.M., Terán-Valdez, A., Siavichay, F.R., Fitzpatrick, S.W. & Coloma, L.A. (2022) Harlequin frog rediscoveries provide insights into species persistence in the face of drastic amphibian declines. Biological Conservation, 276, 109784. https://doi.org/10.1016/j.biocon.2022.109784
  51. Jorge, R.F., Magnusson, W.E., Da Silva, D.A., Polo, É.M. & Lima, A.P. (2020a) Urban growth threatens the lowland Amazonian Manaus harlequin frog which represents an evolutionarily significant unit within the genus Atelopus (Amphibia: Anura: Bufonidae). Journal of Zoological Systematics and Evolutionary Research, 58, 1195–1205. https://doi.org/10.1111/jzs.12390
  52. Jorge, R.F., Ferrão, M. & Lima, A.P. (2020b) 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, 310. https://doi.org/10.3390/d12080310
  53. Jorge, R.F., Lima, A.P. & Stow, A.J. (2022) Selection and localised genetic structure in the threatened Manauense harlequin frog (Bufonidae: Atelopus manauensis). Conservation Genetics, 23, 559–574. https://doi.org/10.1007/s10592-022-01436-8
  54. Kalyaanamoorthy, S., Minh, B.Q., Wong, T.K.F., von Haeseler, A. & Jermiin, L.S. (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods, 14, 587–589. https://doi.org/10.1038/nmeth.4285
  55. Kok, P.J.R., Ratz, S., MacCulloch, R.D., Lathrop, A., Dezfoulian, R., Aubret, F. & Means, D.B. (2018) Historical biogeography of the palaeoendemic toad genus Oreophrynella (Amphibia: Bufonidae) sheds a new light on the origin of the Pantepui endemic terrestrial biota. Journal of Biogeography, 45, 26–36. https://doi.org/10.1111/jbi.13093
  56. Köhler, J., Jansen, M., Rodríguez, A., Kok, P.J.R., Toledo, L.F., Emmrich, M., Glaw, F., Haddad, C.F.B., Rödel, M.-O. & Vences, M. (2017) The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa, 4251 (1), 1–124. https://doi.org/10.11646/zootaxa.4251.1.1
  57. Kimura, M. (1980) A simple method for estimating evolutionary rate of base substitutions through comparative studies of nucleotide sequences. Journal of Molecular Evolution, 16, 111–120. https://doi.org/10.1007/BF01731581
  58. Kumar, S., Stecher, G., Li, M., Knyaz, C. & Tamura, K. (2018) MEGA X: molecular evolutionary genetics analysis across computing platforms. Molecular Biology and Evolution, 35, 1547–1549. https://doi.org/10.1093/molbev/msy096
  59. Langer, M., Boistel, R., Pagot, E., Cloetens, P. & Peyrin, F. (2010) X-ray in-line phase microtomography for biomedical applications. Microscopy: Science, Technology, Applications and Education, 1, 391–402.
  60. La Marca, E. (1983) A new frog of the genus Atelopus (Anura: Bufonidae) from a Venezuelan cloud forest. Contributions to Biology and Geology of the Milwaukee Public Museum, 54, 1–12.
  61. La Marca, E., Lips, K.R., Lötters, S., Puschendorf, R., Ibáñez, R., Rueda-Almonacid, J.V., Schulte, R., Marty, C., Castro, F., Manzanilla-Puppo, J., García-Pérez, J.E., Bolaños, F., Chaves, G., Pounds, J.A., Toral, E. & Young, B.E. (2005) Catastrophic population declines and extinctions in Neotropical harlequin frogs (Bufonidae: Atelopus). Biotropica, 37, 190–201. https://doi.org/10.1111/j.1744-7429.2005.00026.x
  62. Lecocq, T., Vereecken, N.J., Michez, D., Dellicour, S., Lhomme, P., Valterová, I., Rasplus, J.-Y. & Rasmont, P. (2013) Patterns of genetic and reproductive traits differentiation in mainland vs. Corsican populations of bumblebees. PLoS ONE, 8, e65642. https://doi.org/10.1371/journal.pone.0065642
  63. Lescure, J. (“1973” 1974) Présence d’une sous-espéce d’ Atelopus pulcher (Amphibien, Anoure) dans les Guyanes: Atelopus pulcher hoogmoedi. Bulletin du Muséum National d’Histoire Naturelle, 3, 997–1005.
  64. Lescure, J. (“1973” 1974) Contribution à l’étude des amphibiens de Guyane Française. 1. Notes sur Atelopus flavescens Duméril et Bibron et description d’une nouvelle espèce. Vie et Milieu. Série C, Biologie Terrestre, 23, 125–141.
  65. Lescure, J. (1981) Contribution à l’étude des amphibiens de Guyane française. VIII. Validation d’ Atelopus spumarius Cope, 1871, et désignation d’un néotype. Description d’ Atelopus spumarius barbotini nov. ssp. Données étho-écologiques et biogéographiques sur les Atelopus du groupe flavescens (Anoures, Bufonidés). Bulletin du Muséum National d’Histoire Naturelle, 4, 893–910.
  66. Lescure, J. & Gasc, J.P. (1986) Partage de l’espace forestier par les amphibiens et les reptiles en Amazonie du nord-ouest. Caldasia, 15, 707–723.
  67. Lips, K.R., Diffendorfer, J., Mendelson III, J.R. & Sears, M.W. (2008) Riding the wave: reconciling the roles of disease and climate change in amphibian declines. PLoS Biology, 6, e72. https://doi.org/10.1371/journal.pbio.0060072
  68. Lötters, S. (1996) The Neotropical Toad Genus Atelopus. Checklist—Biology—Distribution. M. Vences & F. Glaw, Cologne. 143 pp.
  69. Lötters, S. & De la Riva, I. (1998) Redescription of Atelopus tricolor Boulenger from southeastern Peru and adjacent Bolivia, with comments on related forms. Journal of Herpetology, 32, 481–488. https://doi.org/10.2307/1565201
  70. Lötters, S., Haas, W., Schick, S. & Böhme, W. (2002a) On the systematics of the harlequin frogs (Amphibia: Bufonidae: Atelopus) from Amazonia. II: Redescription of Atelopus pulcher (Boulenger, 1882) from the eastern Andean versant in Peru. Salamandra, 38, 165–184.
  71. Lötters, S., Haas, W., Schick, S. & Böhme, W. (2002b) On the systematics of the harlequin frogs (Amphibia: Bufonidae: Atelopus) from Amazonia. I: Description of a new species from the Cordillera Azul, Peru. Salamandra, 38, 95–104.
  72. Lötters, S. (2003) On the systematics of harlequin frogs (Amphibia: Bufonidae: Atelopus) from Amazonia. III: A new, remarkably dimorphic species from the Cordillera Azul, Peru. Salamandra, 39, 169–180.
  73. Lötters, S., La Marca, E. & Vences, M. (2004) Redescriptions of two toad species of the genus Atelopus from coastal Venezuela. Copeia, 2004, 221–233.
  74. Lötters, S., Schulte, R., Córdova, J.H. & Veith, M. (2005) Conservation priorities for harlequin frogs (Atelopus spp.) of Peru. Oryx, 39, 343–346.
  75. Lötters, S., van der Meijden, A., Rödder, D., Koester, T.E., Kraus, T., La Marca, E., Haddad, C.F.B. & Veith, M. (2010) Reinforcing and expanding the predictions of the disturbance vicariance hypothesis in Amazonian harlequin frogs: a molecular phylogenetic and climate envelope modelling approach. Biodiversity and Conservation, 19, 2125–2146. https://doi.org/10.1007/s10531-010-9869-y
  76. Lötters, S., van der Meijden, A., Coloma, L.A., Boistel, R., Cloetens, P., Ernst, R., Lehr, E. & Veith, M. (2011) Assessing the molecular phylogeny of a near extinct group of vertebrates: the Neotropical harlequin frogs (Bufonidae; Atelopus). Systematics and Biodiversity, 9, 45–57. https://doi.org/10.1080/14772000.2011.557403
  77. 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. https://doi.org/10.3897/herpetozoa.32.e39192
  78. Lötters, S., Plewnia, A., Hönig, A., Jung, A., Laudor, J. & Ziegler, T. (2022) The gastromyzophorous tadpole of the pink harlequin frog from Suriname with comments on the taxonomy of Guianan clade Atelopus (Amphibia, Bufonidae). Zootaxa, 5087 (4), 591–598. https://doi.org/10.11646/zootaxa.5087.4.7
  79. Lötters, S., Plewnia, A., Böning, P., Jung, A., Pinto Erazo, M.A., Velásquez-Trujillo, D.A., Veith, M., La Marca, E., Catenazzi, A., Shepack, A., Neam, K., Valencia, L.M., Acosta-Galvis, A.R., Alarcon Vela, Y., Allen, J.P., Alfaro Segundo, J.O., Almendáriz Cabezas, A. de L., Alvarado Barboza, G., Alves-Silva, K.R., Anganoy-Criollo, M., Rodrigues, M.T., Arbeláez Ortiz, E., Arpi Lojano, J.D., Siavichay Pesántez, F.R., Arteaga, A., Ballestas, O., García Méndez, C.Z., Lampo, M., Barrera Moscoso, D., Herrera-Alva, V., Torres-Ccasani, G., Barros-Castañeda, J.D., Pérez-González, J.L., Rocha-Usuga, A., Rojas Montaño, S., Rueda Solano, L.A., Villalba-Fuentes, J., Batista, A., Quiroz-Espinoza, M., Bernal, M.H., Betancourt, E., López-Perilla, Y.R., Medina-Rangel, G.F., Rivera-Correa, M., da Cunha Bitar, Y.O., Bravo-Valencia, L., Cáceres Andrade, J.F., Cadenas, D., Chaparro Auza, J.C., Chaves-Portilla, G.A., Chávez, G., Roca-Rey Ross, B., Coloma, L.A., Quezada Riera, A.B., Terán-Valdez, A., Cortez-Fernandez, C.F., Courtois, E.A., Culebras, J., De la Riva, I., Díaz, V., Elizondo Lara, L.C., Ernst, R., Flechas, S.V., Foch, T., Fouquet, A., García-Pérez, J.E., Gómez-Hoyos, D.A., Gomides, S.C., Guerrel, J., Gratwicke, B., Guayasamin, J.M., Griffith, E., Ibáñez, R., Idrovo, C.I., Jiménez Monge, A., Jorge, R.F., Klocke, B., Lehr, E., Lewis, C.H.R., Lindquist, E.D., Mazepa, G., Merino Viteri, A., Mulder, K., Pacheco-Suarez, M., Pereira-Muñoz, A., Tovar-Siso, P.C., Pisso Florez, A.G., Ponce, M., Poole, V., Quiroz, A.J., Ramírez Guerra, A., Ramírez, J.P., Reichle, S., Reizine, H., Rößler, D.C., Señaris, C., Sorokin, A., Venegas, P.J., von May, R. & Webster Bernal, J.F. (2023) Ongoing harlequin toad declines suggest the amphibian extinction crisis is still an emergency. Communications Earth and Environment, 4, 412. https://doi.org/10.1038/s43247-023-01069-w
  80. Luedtke, J.A., Chanson, J., Neam, K ., Hobin L., Maciel, A.O., Catenazzi, A., Borzée, A., Hamidy, A., Aowphol, A., Jean, A., Sosa-Bartuano, A., Fong G., A., de Silva, A., Fouquet, A., Angulo, A., Kidov, A.A., Muñoz Saravia, A., Diesmos, A.C., Tominaga, A., Shrestha, A., Gratwicke, B., Tjaturadi, B., Martínez Rivera, C.C., Vásquez Almazán, C.R., Señaris, C., Chandramouli, S.R., Strüssmann, C., Cortez Fernández, C.F., Azat, C., Hoskin, C.J., Craig Hilton-Taylor, C., Whyte, D.L., Gower, D.J., Olson, D.H., Cisneros-Heredia, D.F., Santana, D.J., Nagombi, E., Najafi-Majd, E., Quah, E.S.H., Bolaños, F., Xie, F., Brusquetti, F., Álvarez, F.S., Andreone, F., Glaw, F., Castañeda, F.E., Kraus, F., Parra-Olea, G., Chaves, G., Medina-Rangel, G.F., González-Durán, G., Ortega-Andrade, H.M., Machado, I.F., Das, I., Ribeiro Dias, I., Urbina-Cardona, J.N., Crnobrnja-Isailović, J., Yang, J.H., Jianping, J., Wangyal, J.T., Rowley, J.J.L., Measey, J., Vasudevan, K., Chan, K.O., Gururaja, K.V., Ovaska, K., Warr, L.C., Canseco-Márquez, L., Toledo, L.F., Díaz, L.M., Khan, M.M.H., Meegaskumbura, M., Acevedo, M.E., Napoli, M.F., Ponce, M.A., Vaira, M., Lampo, M., Yánez-Muñoz, M.H., Scherz, M.D., Rödel, M.O., Matsui, M., Fildor, M., Kusrini, M.D., Ahmed, M.F., Rais, M., Kouamé, N.G., García, N., Gonwouo, N.L., Burrowes, P.A., Imbun, P.Y., Wagner, P., Kok, P.J.R., Joglar, R.L., Auguste, R.J., Albuquerque Brandão, R., Ibáñez, R., von May, R., Hedges, S.B., Biju, S.D., Ganesh, S.R., Wren, S., Das, S., Flechas, S.V., Ashpole, S.L., Robleto-Hernández, S.J., Loader, S.P., Incháustegui, S.J., Garg, S., Phimmachak, S., Richards, S.J., Slimani, T., Osborne-Naikatini, T., Abreu-Jardim, T.P.F., Condez, T.H., De Carvalho, T.R., Cutajar, T.P., Pierson, T.W., Nguyen, T.Q., Kaya, U., Yuan, Z., Long, B., Langhammer, P. & Stuart, S.N. (2023) Ongoing declines for the world’s amphibians in the face of emerging threats. Nature, 622, 308–314. https://doi.org/10.1038/s41586-023-06578-4
  81. Lynch, J.D. (1986) Notes on the reproductive biology of Atelopus subornatus. Journal of Herpetology, 20, 126–129.
  82. Lynch, J.D. (1993) A new harlequin frog from the Cordillera Oriental of Colombia (Anura, Bufonidae, Atelopus). Alytes, 11, 77–87.
  83. Marcillo-Lara, A., Coloma, L.A., Álvarez-Solas, S. & Terneus, E. (2020) The gastromyzophorous tadpoles of Atelopus elegans and A. palmatus (Anura: Bufonidae), with comments on oral and suction structures. Neotropical Biodiversity, 6, 1–13. https://doi.org/10.1080/23766808.2019.1709378
  84. McDiarmid, R.W. (1971) Comparative morphology and evolution of frogs of the Neotropical genera Atelopus, Dendrophryniscus, Melanophryniscus and Oreophrynella. Bulletin of the Los Angeles County Museum of Natural History, 12, 1–66.
  85. McDiarmid, R.W. (1973) A new species of Atelopus (Anura, Bufonidae) from northeastern South America. Contributions in Science. Natural History Museum of Los Angeles County, 240, 1–12.
  86. Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M. D., van Haeseler, A. & Lanfear R. (2020) IQ-TREE 2: New models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution, 37, 1530–1534. https://doi.org/10.1093/molbev/msaa015
  87. Miyata, K. (1980) A new species of Atelopus (Anura: Bufonidae) from the cloud forests of northwestern Ecuador. Breviora, 458, 1–10.
  88. Moraes, L.J.C.L., Werneck, F.P., Réjaud, A., Rodrigues, M.T., Prates, I., Glaw, F., Kok, P.J.R., Ron, S.R., Chaparro, J.C., Osorno-Muñoz, M., Dal Vechio, F., Recoder, R.S., Marques-Souza, S., Rojas, R.R., Demay, L., Hrbek, T. & Fouquet, A. (2022) Diversification of tiny toads (Bufonidae: Amazophrynella) sheds light on ancient landscape dynamism in Amazonia. Biological Journal of the Linnean Society, 136, 75–91. https://doi.org/10.1093/biolinnean/blac006
  89. Noonan, B.P. & Gaucher, P. (2005) Phylogeography and demography of Guianan harlequin toads (Atelopus): diversification within a refuge. Molecular Ecology, 14, 3017–3031. https://doi.org/10.1111/j.1365-294X.2005.02624.x
  90. Osorno-Muñoz, M., Ardila-Robayo, M.C. & Ruíz-Carranza, P. (2001) Tres nuevas especies del género Atelopus A.M.C. Dumeril & Bibron 1841 (Amphibia: Bufonidae) de las partes altas de la Cordillera Oriental Colombiana. Caldasia, 23, 509–522.
  91. Padial, J.M., A. Miralles, I. De la Riva & M. Vences (2010) The integrative future of taxonomy. Frontiers in Zoology, 7, 16. https://doi.org/10.1186/1742-9994-7-16
  92. Patzelt, E. (1989) Fauna del Ecuador. Banco Central del Ecuador, Quito, 433 pp.
  93. Pearson, K.C. & Tarvin, R.D. (2022) A review of chemical defense in harlequin toads (Bufonidae: Atelopus). Toxicon, 13, 100092.
  94. Pérez-Gonzalez, J.L., Rada, M., Vargas-Salinas, F. & Rueda-Solano, L.A. (2020) The tadpoles of two Atelopus species (Anura: Bufonidae) from the Sierra Nevada de Santa Marta, Colombia, with notes on their ecology and comments on the morphology of Atelopus larvae. South American Journal of Herpetology, 15, 47–62.
  95. Peters, J.A. (1973) The frog genus Atelopus in Ecuador. Smithsonian Contributions to Zoology, 145, 1–49.
  96. Plewnia, A., Lötters, S., Gomides, S., De Agrò, M. & Rößler, D.C. (2024a) Sex-specific ventral dichromatism and melanization in harlequin toads (Atelopus): a common but overlooked character of unknown function. Evolutionary Ecology, 38, 571–583. https://doi.org/10.1007/s10682-024-10288-2
  97. Plewnia, A., Terán-Valdez, A., Culebras, J., Boistel, R., Paluh, D.J., Quezada Riera, A.B., Heine, C.H., Reyes-Puig, J.P., Salazar-Valenzuela, D., Guayasamin, J.M. & Lötters, S. (2024b) A new species of harlequin toad (Bufonidae: Atelopus) from Amazonian Ecuador. Salamandra, 60, 237–253.
  98. Pons, J., Barraclough, T.G., Gomez-Zurita, J., Cardoso, A., Duran, D. P., Hazell, S., Kamoun S., Sumlin, W.D. & Vogler, A.P. (2006) Sequence-based species delimitation for the DNA taxonomy of undescribed insects. Systematic Biology, 55, 595–609. https://doi.org/10.1080/10635150600852011
  99. Portik, D.M., Streicher, J. & Wiens, J.J. (2023) Frog phylogeny: A time-calibrated, species-level tree based on hundreds of loci and 5,242 species. Molecular Phylogenetics and Evolution, 188, 107907. https://doi.org/10.1016/j.ympev.2023.107907
  100. Pramuk, J.B. (2006) Phylogeny of South American Bufo (Anura: Bufonidae) inferred from combined evidence. Zoological Journal of the Linnean Society, 146, 407–452. https://doi.org/10.1111/j.1096-3642.2006.00212.x
  101. Puillandre, N., Brouillet, S. & Achaz, G. (2020) ASAP: Assemble Species by Automatic Partitioning. Molecular Ecolology Resources, 21, 609–620. https://doi.org/10.1111/1755-0998.13281
  102. Rambaut, A., Drummond, A.J., Xie, D., Baele, G. & Suchard, M.A. (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology, 67, 901–904. https://doi.org/10.1093/sysbio/syy032
  103. Ramírez, J., Jaramillo, C., Lindquist, E., Crawford, A. & Ibáñez, R. (2020) Recent and rapid radiation of the highly endangered harlequin frogs (Atelopus) into Central America inferred from mitochondrial DNA sequences. Diversity, 12, 360. https://doi.org/10.3390/d12090360
  104. Read, M. (2018) Monitoreo biológico Yasuní. Anfibios y reptiles. Vol. 4. Ecuambiente Consulting Group, Quito, 248 pp.
  105. Ringler, M., Himmel, T., Werner, P., Luger, M., Rößler, D.C., Ringler, E. & Lötters, S. (2022) Male life history of a harlequin toad population in French Guiana. Salamandra, 58, 171–186.
  106. Rivero, J.A. (1963) Five new species of Atelopus from Colombia, with notes on other forms from Colombia and Ecuador. Caribbean Journal of Science, 3, 103–124.
  107. Rivero, J.A. (1968) More on the Atelopus (Amphibia, Salientia) from western South America. Caribbean Journal of Science, 8, 19–28.
  108. Rivero, J.A. & Granados Díaz, H. (1993) Nueva especie de Atelopus (Amphibia: Bufonidae) del Departamento del Cauca, Colombia. Caribbean Journal of Science, 29, 12–17.
  109. Roček, Z., Přikryl, T., Baleeva, N., Vazeille, A., Boistel, R., Bravin, A., Nemoz, C., van Dijk, E., Smirina, E.M. & Claessens, L. (2016) Contribution to the head anatomy of the basal frog Barbourula busuangensis and the evolution of the Anura. Russian Journal of Herpetology, 23, 163–194.
  110. Rocha Usuga, A.A., Vargas Salinas, F. & Rueda-Solano, L.A. (2017) Not every drought is bad: Quantifying reproductive effort in the harlequin frog Atelopus laetissimus (Anura: Bufonidae). Journal of Natural History, 51, 1913–1928. https://doi.org/10.1080/00222933.2017.1355075
  111. Rößler, D.C., Lötters, S., Mappes, J., Valkonen, J.K., Menin, M., Lima, A.P. & Pröhl, H. (2019) Sole coloration as an unusual aposematic signal in a Neotropical toad. Scientific Reports, 9, 1128. https://doi.org/10.1038/s41598-018-37705-1
  112. Ron, S., Bustamante, M., Coloma, L.A. & Mena, B. (2009) Sapos. Museo de Zoología, Pontificia Universidad Católica del Ecuador, Balsa de Sapos, Fundación Otonga, Amphibian Ark, Quito, 256 pp.
  113. Ronquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19, 1572–1574. https://doi.org/10.1093/sysbio/46.1.195
  114. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. https://doi.org/10.1093/sysbio/sys029
  115. Rueda-Almonacid, J.V., Rodríguez-Mahecha, J.V., La Marca, E., Lötters, S., Kahn, T. & Angulo, A. (Eds.) (2005) Ranas Arlequines. Serie Libretas de Campo. 5. Conservación Internacional Colombia, Bogotá, 158 pp.
  116. Rueda-Solano, L.A. (2012) Atelopus carrikeri Ruthven, 1916 (Anura: Bufonidae) una especie policromatica. Herpetotropicos, 8, 61–66.
  117. Rueda-Solano, L.A., Vargas Salinas, F., Gonzalez Pérez, J.I., Sanchez A., Ramírez, A., Navas C.A. & Crawford A.J. (2022) Mate-guarding behavior in anurans: intra-sexual selection and the evolution of prolonged amplexus in the harlequin toad Atelopus laetissimus. Animal Behaviour, 185, 127–142. https://doi.org/10.1016/j.anbehav.2021.12.003
  118. Ruíz-Carranza, P.M. & Hernández-Camacho, J.I. (1978) Una nueva especie Colombiano de Atelopus (Amphibia: Bufonidae). Caldasia, 12, 181–197.
  119. Ruíz-Carranza, P.M. & Osorno-Muñoz, M. (1994) Tres nuevas especies de Atelopus A.M.C. Duméril & Bibron 1841 (Amphibia: Bufonidae) de la Cordillera Central de Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 19, 165–179.
  120. Savage, J.M. (1972) The harlequin frogs, genus Atelopus, of Costa Rica and western Panama. Herpetologica, 28, 77–94.
  121. Savage, J.M. & Heyer, R.W. (1997) Digital webbing formulae for anurans: a refinement. Herpetological Review, 28, 131.
  122. Sexton, O. (1958) Observations of the life history of a Venezuelan frog, Atelopus cruciger. Acta Biologica Venezuelica, 2, 235–242.
  123. Standish, I., Leis, E., Schmitz, N., Credico, J., Erickson, S., Bailey, J., Kerby, J., Phillips, K. & Lewis, T. (2018) Optimizing, validating, and field testing a multiplex qPCR for the detection of amphibian pathogens. Diseases of Aquatic Organisms, 129, 1–13.
  124. Sueur, J., Aubin, T. & Simonis, C. (2008) Seewave: a free modular tool for sound analysis and synthesis. Bioacoustics, 18, 213–226.
  125. Tarvin, R.D., Peña, P. & Ron, S.R. (2014) Changes in population size and survival in Atelopus spumarius (Anura: Bufonidae) are not correlated with chytrid prevalence. Journal of Herpetology, 48, 291–197.
  126. Valencia, L.M. & Marin da Fonte, L.F. (2022) Collaborative work brings hope for threatened harlequin toads. Oryx, 56, 12.
  127. Van Bocxlaer, I., Loader, S.P., Roelants, K., Biju, S.D., Menegon, M. & Bossuyt, F. (2010) Gradual adaptation toward a range-expansion phenotype initiated the global radiation of toads. Science, 327, 679–682. https://doi.org/10.1126/science.1181707
  128. Vélez-Rodríguez, C.M. & Ruíz-Carranza, P.M. (1997) Una nueva especie de Atelopus (Amphibia: Anura: Bufonidae) de la Cordillera Central, Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Naturales, 21, 555–563.
  129. Vences, M., Miralles, A., Brouillet, S., Ducasse, J., Fedosov, A., Kharchev, V., Kostadinov, I., Kumari, S., Patmanidis, S., Scherz, M.D., Puillandre, N. & Renner, S.S. (2021) iTaxoTools 0.1: Kickstarting a specimen-based software toolkit for taxonomists. Megataxa, 6, 77–92. https://doi.org/10.11646/megataxa.6.2.1
  130. Venegas, P.J., Catenazzi, A., Siu-Ting, K. & Carrillo, J. (2008) Two new harlequin frogs (Anura: Atelopus) from the Andes of northern Peru. Salamandra, 44, 163–176.
  131. Veselý, M. & Batista, A. (2021) A new species of Atelopus (Amphibia: Bufonidae) from eastern Panama. Zoological Research, 42, 272–279. https://doi.org/10.24272/j.issn.2095-8137.2020.319
  132. Weitkamp, T., Scheel, M., Perrin, J., Daniel, G., King, A., Le Roux, V., Giorgetta, J.-L., Carcy, A., Langlois, F., Desjardins, K., Menneglier, C., Cerato, M., Engblom, C., Cauchon, G., Moreno, T., Rivard, C., Gohon, Y. & Polack, F. (2022) Microtomography on the ANATOMIX beamline at Synchrotron SOLEIL. Journal of Physics: Conference Series, 2380, 012122. https://doi.org/10.1088/1742-6596/2380/1/012122
  133. Wiley, E.O. (1978) The evolutionary species concept reconsidered. Systematic Biology, 27 (1), 17–26. https://doi.org/10.2307/2412809
  134. Wiley, E.O. & Liebermann, B.S. (2011) Phylogenetics: The Theory and Practice of Phylogenetic Systematics. 2nd Edition. Wiley-Blackwell, Hoboken, New Jersey, 406 pp.
  135. Womack, M.C., Christensen-Dalsgaard, J., Coloma, L.A. & Hoke, K.L. (2018) Sensitive high-frequency hearing in earless and partially eared harlequin frogs (Atelopus). Journal of Experimental Biology, 221, jeb169664. https://doi:10.1242/jeb.169664
  136. Zanette, I., Daghfous, G., Weitkamp, T., Gillet, B., Adriaens, D., Langer, M., Cloetens, P., Helfen, L., Bravin, A., Peyrin, F., Baumbach, T., Dischler, J.-M., Van Loo, D., Praet, T., Poirier-Quinot, M. & Boistel, R. (2014) Looking inside marine organisms with magnetic resonance and X‐ray imaging. In: Reynaud, E.G. (Ed.), Imaging Marine Life: Macrophotography and Microscopy Approaches for Marine Biology. Wiley-Blackwell, Hoboken, New Jersey, pp. 122–184.