Abstract
The genus Speonemadus Jeannel, 1922 currently includes 12 species distributed in the Western Mediterranean (Iberian Peninsula, North Africa, Sicily and continental Italy). Two new species are described from Morocco, S. brusteli sp. n. from the Rif and S. comasi sp. n. from the High Atlas, the first in the S. vandalitiae-group of species and the second with an isolated position within the genus. Two species are also resurrected from synonymy, S. pulchellus (Reitter, 1885) stat. rest. (former synonym of S. orchesioides (Fairmaire, 1879)) and S. gracilis (Kraatz, 1870) stat. rest. (former synonym of S. vandalitiae (Heyden, 1870)), raising the total number of species of the genus to 16. Lectotypes are designated for Anemadus pulchellus Reitter, 1885, and Anemadus tenuipes Peyerimhoff, 1917. A molecular phylogeny of the genus Speonemadus is presented, based on a combination of mitochondrial and nuclear genes and including seven of the currently recognised species, one of the newly described (S. brusteli sp. n.) and the two reinstated taxa. The genus is estimated to have diversified from the late Miocene (Tortonian), with successive cladogenetic events related to the isolation of landmasses between Iberia and North Africa, including the opening of the Strait of Gibraltar at the end of the Messinian. The distribution of some of the species of Speonemadus remains still uncertain, due to their frequent misidentification and the problems with their taxonomic status.
References
Agirre, E. & Blas, M. (2009) Estudio de los Cholevinae (Coleoptera: Leiodidae) de las Sierras de Ancares y del Caurel (Noroeste de la Península Ibérica). Elytron, 22–23 (2008–2009), 87–98.
Andújar, C., Gómez-Zurita, J., Rasplus, J.-Y. & Serrano, J. (2012) Molecular systematics and evolution of the subgenus Mesocarabus Thomson, 1875 (Coleoptera: Carabidae: Carabus), based on mitochondrial and nuclear DNA. Zoological Journal of the Linnean Society, 166, 787–804.
https://doi.org/10.1111/j.1096-3642.2012.00866.x
Antunes-Carvalho, C., Ribera, I., Beutel, R.G. & Gnaspini, P. (2018) Morphology-based phylogenetic reconstruction of Cholevinae (Coleoptera: Leiodidae): a new view on higher-level relationships. Cladistics. [published online]
https://doi.org/10.1111/cla.12230Arano, B., Llorente, G.A., Montori, A., Buckley, D. & Herrero, P. (1998) Diversification in North-West African water frogs: molecular and morphological evidence. Herpetological Journal, 8, 57–64.
Barraclough, T.G., Hogan, J.E. & Vogler, A.P. (1999) Testing whether ecological factors promote cladogenesis in a group of tiger beetles (Coleoptera: Cicindelidae). Proceedings of the Royal Society B, Biological Sciences, 266, 1061–1067.
https://doi.org/10.1098/rspb.1999.0744
Besson, D. (2005) Architecture du bassin rhodano-provencal miocène (Alpes, SE France): relations entre déformation, physiographie et sédimentation dans un bassin molassique d'avant-pays. Paris, École Nationale Supérieure des Mines de Paris, 438 pp.
Beukema, W., De Pous, Ph., Donaire, D., Escoriza, D., Bogaerts, S., Toxopeus, A.G., De Bie, C.A.J.M., Roca, J. & Carranza, S. (2010) Biogeography and contemporary climatic differentiation among Moroccan Salamandra algira. Biological Journal of the Linnean Society, 101, 626–641.
https://doi.org/10.1111/j.1095-8312.2010.01506.x
Blas, M. (1976) Coleópteros cavernícolas del distrito andaluz (Catopidae). Actas 4 Congreso Nacional de Espeleología. Nerja, Málaga, 1976, 157–160.
Blas, M. (1977) Contribución al conocimiento de los Anemadinae de la Península Ibérica. (Col. Catopidae). Comunicacions 6è Simposium d'Espeleologia: Bioespeleologia, Terrassa, 1977, 125–130.
Blas, M. (1979) Contribución al conocimiento de los Catopidae (excepto la subfam. Bathysciinae) de la Península Ibérica (Coleoptera Staphylinoidea). Unpublished PhD thesis, Universidad de Barcelona, Barcelona, 361 pp.
Blas, M. (1981a) Sobre la presencia de Hormosacus vandalitiae (Heyden, 1870) en la isla de Mallorca. (Col. Catopidae). Rapport Communications Internationales Mer Méditerranée, 27 (9), 87–88.
Blas, M. (1981b) La subfamilia Anemadinae Jeannel (Col. Catopidae) i llur distribució geográfica actual. Treballs de l'Institució Catalana d'Història Natural, 9, 155–156.
Blas, M. (1984) El género Hormosacus Jeannel, 1936 y su distribución geográfica. (Col. Catopidae). Mémoires de Biospéologie, 11, 251–255.
Blas, M. (1985) Sobre los Catopidae de Portugal (Coleoptera). Suplemento 1. Boletim da Sociedade Portuguesa de Entomologia, 1985, 169–176.
Blas, M. (1992) Nuevos datos sobre los Cholevidae (Col.) de las Islas Baleares. Elytron, 6, 159–162.
Cieslak, A., Fresneda, J. & Ribera, I. (2014) Life-history specialization was not an evolutionary dead-end in Pyrenean cave beetles. Proceedings of the Royal Society B, Biological Sciences, 281, 20132978.
https://doi.org/10.1098/rspb.2013.2978Cobos, A. (1954) Una breve campaña entomológica por las sierras de Tejeda y Almijara (Provincia de Málaga): Ins. Coleópteros. Archivos del Instituto de Aclimatación de Almería, 2, 81–88.
Coiffait, H. (1954) Catopidés récoltés en Andalousie. Description de trois formes nouvelles. Notes Biospéologiques, 9 (1), 21–24.
Dowsett, H., Barron, J.A., Poore, R.Z., Thompson, R.S., Cronin, T.M., Ishman, S.E. & Willard, D.A. (1999) Middle Pliocene paleoenvironmental reconstruction: PRISM2. U.S. Geological Survey Open File Report 99–535.
https://doi.org/10.3133/ofr99535
Drummond, A.J. & Rambaut, A. (2007) BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology, 7, 214.
https://doi.org/10.1186/1471-2148-7-214
Fabbri, R.A. & Zoia, S. (1996) Segnalazioni faunistiche n° 10—Speonemadus orchesioides (Fairmaire, 1879) (Insecta, Coleoptera, Cholevidae). Quaderni di Studi e Notizie di Storia Naturale della Romagna, 6, 78.
Faille, A., Andújar, C., Fadrique, F. & Ribera, I. (2014) Late Miocene origin of an Ibero-Maghrebian clade of ground beetles with multiple colonizations of the subterranean environment. Journal of Biogeography, 41, 1979–1990.
https://doi.org/10.1111/jbi.12349
Faille, A., Bourdeau, C. & Fresneda, J. (2010b) A new species of blind Trechinae from the Pyrenees of Huesca, and its position within Aphaenops (sensu stricto) (Coleoptera: Carabidae: Trechini). Zootaxa, 2566, 49–56.
Faille, A., Ribera, I., Deharveng, L., Bourdeau, C., Garnery, L., Quéinnec, E. & Deuve, T. (2010a) A molecular phylogeny shows the single origin of the Pyrenean subterranean Trechini ground beetles (Coleoptera: Carabidae). Molecular Phylogenetics and Evolution, 54, 97–105.
https://doi.org/10.1016/j.ympev.2009.10.008
Fairmaire, L. (1879) Descriptions de coléoptères nouveaux du nord de l'Afrique. Annales de la Société Entomologique de France, 5ème Série, 9, 155–172.
Faria e Silva, I., Salgado, J.M. & Serrano, A.R.M. (2013) New data on the Cholevidae (Coleoptera) from Portugal. Graellsia, 69, 45–56.
https://doi.org/10.3989/graellsia.2013.v69.078
Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299.
Fresneda, J. (2013) Los leiódidos (Coleoptera, Leiodidae) subterráneos de Jaén In: Pérez Fernández, T. & Pérez Ruiz, A. (Eds.), Los invertebrados de hábitats subterráneos de Jaén. Grupo de Espeleología de Villacarrillo, Jaén, pp. 164–175.
Fresneda, J., Cárdenas, A.M., Castro, A., Lencina, J.L., López-Colón, J.I. & Baena, M. (2007) Nuevos datos de los Cholevidae en la Península Ibérica (Coleoptera). Boletín de la Asociación Española de Entomología, 31, 187–214.
Fresneda, J., Grebennikov, V. & Ribera, I. (2011) The phylogenetic and geographic limits of Leptodirini (Coleoptera: Leiodidae: Cholevinae), with a description of Sciaphyes shestakovi sp.n. from the Russian Far East. Arthropods Systematics & Phylogeny, 69, 99–123.
Fresneda, J. & Salgado, J.M. (2017) Catálogo de los coleópteros Leiodidae Cholevinae Kirby, 1837 de la península Ibérica e islas Baleares. Monografies del Museu de Ciències Naturals, 7 (2016), 1–308.
Fuente, J.M. de la (1925) Catálogo sistemático-geográfico de los coleópteros observados en la península Ibérica, Pirineos propiamente dichos y Baleares. Boletín de la Real Sociedad Entomológica de España, 7 (2–3), 93–117. [1924]
García-Castellanos, D., Estrada, F., Jiménez-Munt, I., Gorini, C., Fernández, M., Vergés, J. & Vicente, R. de (2009) Catastrophic flood of the Mediterranean after the Messinian salinity crisis. Nature, 462, 778–781.
https://doi.org/10.1038/nature08555
Giachino, P.M. (1985) Choleva (s. str.) gabriellae e Sciodrepoides casalei nuove specie della Kabylia e note corologiche sui Catopidi d'Algeria (Coleoptera, Catopidae). Bollettino del Museo Regionale di Scienze Naturali, Torino, 3, 337–348.
Giachino, P.M. & Vailati, D. (1993) Revisione degli Anemadinae. Monografie di "Natura Bresciana" No. 18. Museo Cívico di Scienze Naturali di Brescia, Brescia, 314 pp.
Giachino, P.M. & Vailati, D. (2000) Nuovi dati sugli Anemadinae con descrizione di Anemadus whiteheadi sp. n. (Coleotera Cholevidae). Elytron, 14, 119–124.
Giachino, P.M. & Vailati, D. (2007) I Coleotteri Cholevidi dell’Appennino settentrionale e centrale: inventario, analisi faunistica e origine del popolamento (Coleoptera Cholevidae). Biogeographia, 28, 365–420.
Haywood, A.M., Sellwood, B.W. & Valdes, P.J. (2000) Regional warming: Pliocene (3Ma) paleoclimate of Europe and the Mediterranean. Geology, 28, 1063–1066.
https://doi.org/10.1130/0091-7613(2000)28%3C1063:RWPMPO%3E2.0.CO;2
Hebert, P.D.N., Penton, E.H., Burns, J.M., Janzen, D.H. & Hallwachs, W. (2004) Ten species in one: DNA barcoding reveals cryptic species in the Neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences of the United States of America, 101, 14812–14817.
https://doi.org/10.1073/pnas.0406166101
Heyden, L. von (1870) Catops Vandalitiae Heyd. nov. sp. In: Heyden, L. von, Entomologische Reise nach dem südlichen Spanien, der Sierra Guadarrama und Sierra Morena, Portugal und den Cantabrischen Gebirgen. Mit Beschreibungen der neuen Arten. Nicolaische Verlagsbuchhandlung, Berlin, pp. 97–98.
Heyden, L. von (1881) Zweites Verzeichniss von Coleopteren aus Asturien. Deutsche Entomologische Zeitschrift, 25, 241–246.
Heyden, L. von (1883) Zwei neue Käfer von Creta. Deutsche Entomologische Zeitschrift, 27, 368.
Hidalgo-Galiana, A. & Ribera, I. (2011) Late Miocene diversification of the genus Hydrochus (Coleoptera, Hydrochidae) in the west Mediterranean area. Molecular Phylogenetics and Evolution, 59, 377–385.
https://doi.org/10.1016/j.ympev.2011.01.018
ISC (2017) International Chronostratigraphic Chart. Version 2017/02. International Commission on Stratigraphy. Available from: http://www.stratigraphy.org (accessed 9 May 2018)
Jeannel, R. (1922) Silphidae Catopinae, (deuxième série). Étude phylogénétique et paléogéographique de la sous-famille. Archives de Zoologie Expérimentale et Générale, 61, l–98.
Jeannel, R. (1936) Monographie des Catopidés. Mémoires du Museum National d'Histoire Naturelle, 1, 1–433.
Jolivet, L., Augier, R., Robin, C., Suc, J.-P. & Rouchy, J.M. (2006) Lithospheric-scale geodynamic context of the Messinian salinity crisis. Sedimentary Geology, 188–189, 9–33.
https://doi.org/10.1016/j.sedgeo.2006.02.004
Katoh, K. & Toh, H. (2008) Recent developments in the MAFFT multiple sequence alignment program. Briefings in Bioinformatics, 9, 286–298.
https://doi.org/10.1093/bib/bbn013
Kraatz, G. (1870) Choleva gracilis n. sp. In: Heyden, L. von, Entomologische Reise nach dem südlichen Spanien, der Sierra Guadarrama und Sierra Morena, Portugal und den Cantabrischen Gebirgen. Mit Beschreibungen der neuen Arten. Nicolaische Verlagsbuchhandlung, Berlin, pp. 99–101.
López, H. & Oromí, P. (2010) A pitfall trap for sampling the mesovoid shallow substratum (MSS) fauna. Speleobiology Notes, 2, 7–11.
Martínez de la Escalera, M. (1927) Del verdadero hábitat de algunos catopinos (Col. Silphidae). Boletín de la Real Sociedad Española de Historia Natural, 27, 426–428.
Meulenkamp, J.E. & Sissingh, W. (2003) Tertiary palaeogeography and tectonostratigraphic evolution of the Northern and Southern Peri-Tethys platforms and the intermediate domains of the African-Eurasian convergent plate boundary zone. Palaeogeography, Palaeoclimatology, Palaeoecology, 196, 209–228.
https://doi.org/10.1016/S0031-0182(03)00319-5
Paulo, O.S., Pinheiro, J., Miraldo, A., Bruford, M.W., Jordan, W.C. & Nichols, R.A. (2008) The role of vicariance vs. dispersal in shaping genetic patterns in ocellated lizard species in the western Mediterranean. Molecular Ecology, 17, 1535–1551.
https://doi.org/10.1111/j.1365-294X.2008.03706.x
Pérez, T. (2015) Catálogo de invertebrados citados en cuevas, simas y minas de Andalucía. Monografías Bioespeleológicas, 9, 1–97.
Pérez, T. & Pérez, A. (2006) Estudios bioespeleológicos en la Cueva Secreta del Sagreo, La Iruela (Jaén). Monografías Bioespeleológicas, 1, 1–13.
Pérez, T., Pérez, A., Pérez, J. & García, F. (2013) La cueva Secreta del Sagreo (La Iruela, Jaén, sur de España), una cavidad con historia en Jaén. Espeleo-Tema, 24, 31–39.
Pérez, A. & Tinaut, A. (2005) Bioespeleología en la provincia de Jaén. Espeleo, 17, 1–10.
Perreau, M. (2000) Catalogue des coléoptères Leiodidae Cholevinae et Platypsyllinae. Mémoires de la Société Entomologique de France, 4, 1–460.
Perreau, M. (2004) Leiodidae. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 2. Apollo Books, Stenstrup, pp. 133–203.
Peyerimhoff, P. de (1917) Nouveaux coléoptères du Nord-Africain. Vingt-cinquième note avec des renseignements synonymiques sur les Silphidae et les Histeridae. Annales de la Société Entomologique de France, 86, 117–148.
Reboleira, A.S., Fresneda, J. & Salgado, J.M. (2017) A new species of Speonemadus from Portugal with the revision of the escalerai-group (Coleoptera, Leiodidae). European Journal of Taxonomy, 261, 1–23.
https://doi.org/10.5852/ejt.2017.261
Recuero, E., Iraola, A., Rubio, X., Machordom, A. & García-París, M. (2007) Mitochondrial differentiation and biogeography of Hyla meridionalis (Anura: Hylidae): an unusual phylogeographical pattern. Journal of Biogeography, 34, 1207–1219.
https://doi.org/10.1111/j.1365-2699.2007.01688.x
Reitter, E. (1885) Bestimmungs-Tabellen der europäischen Coleopteren, 12 Necrophaga. Verhandlungen des Naturforschenden Vereines in Brünn, 23 (1884), 3–122.
Ribera, C. (1970) Resultados faunísticos de la campaña Serranía de Ronda 70. Actas I Congreso Nacional de Espeleología, Barcelona, 123–129.
Ribera, I., Fresneda, J., Bucur, R., Izquierdo, A., Vogler, A.P., Salgado, J.M. & Cieslak, A. (2010) Ancient origin of a western Mediterranean radiation of subterranean beetles. BMC Evolutionary Biology, 10, 29.
https://doi.org/10.1186/1471-2148-10-29
Rosenbaum, G., Lister, G.S. & Duboz, C. (2002) Reconstruction of the tectonic evolution of the western Mediterranean since the Oligocene. Journal of the Virtual Explorer, 8, 107–130.
https://doi.org/10.3809/jvirtex.2002.00053
Rubink, W.L., Murray, K.D., Baum, K.A. & Pinto, M.A. (2003) Long term preservation of DNA from honey bees (Apis mellifera) collected in aerial pitfall traps. Texas Journal of Science, 55, 159–168.
Sáez Bolaño, J. & Blanco Villero, J.M. (2010) Los colévidos epigeos (Coleoptera, Leiodidae, Cholevinae) de la Sierra de Tudía (Badajoz, Extremadura, España). Boletín de la Sociedad Entomológica Aragonesa, 47, 357–361.
Salgado, J.M. (1985) Nuevos datos sobre Catopidae (Col.) capturados en cuevas. Boletín de la Asociación Española de Entomología, 9, 271–277.
Salgado, J.M. (1996) Leiodidae Cholevinae (Coleoptera) in the Orocantabrian phytogeographic province (Cantabrian Cordillera, North Spain). Giornale Italiano di Entomologia, 8, 213–229.
Salgado, J.M., Blas, M. & Fresneda, J. (2008) Coleoptera, Cholevidae. Fauna Ibérica. Vol. 31. Museo Nacional de Ciencias Naturales, CSIC, Madrid, 799 pp.
Salgado, J.M. & Fernández, M. (1998) Estudio de los Leiodidae: Cholevinae (Coleoptera) en las cuencas de los ríos Bernesga, Torío y Porma (León, España). Boletín de la Asociación Española de Entomología, 22, 81–97.
Schettino, A. & Turco, E. (2006) Plate kinematics of the Western Mediterranean region during the Oligocene and Early Miocene. Geophysical Journal International, 166, 1398–1423.
https://doi.org/10.1111/j.1365-246X.2006.02997.x
Seabra, A.F. de (1943) Contribuçoes para o inventário da fauna lusitânica, Insecta, Coleoptera. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 142, 1–151.
Shull, V.L., Vogler, A.P., Baker, M.D., Maddison, D.R. & Hammond, P.M. (2001) Sequence alignment of 18S ribosomal RNA and the basal relationships of adephagan beetles: evidence for monophyly of aquatic families and the placement of Trachypachidae. Systematic Biology, 50, 945–969.
https://doi.org/10.1080/106351501753462894
Simon, C., Frati, F., Beckenbach, A., Crespi, B., Liu, H. & Flook, P. (1994) Evolution, weighting, and phylogenetic utility of mitochondrial gene-sequences and a compilation of conserved polymerase chain-reaction primers. Annals of the Entomological Society of America, 87, 651–701.
https://doi.org/10.1093/aesa/87.6.651
Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology, 57, 758–771.
https://doi.org/10.1080/10635150802429642
Suc, J.-P. (1984) Origin and evolution of the Mediterranean vegetation and climate in Europe. Nature, 307, 429–432.
https://doi.org/10.1038/307429a0
Szymczakowski, W. (1970) Contribution à la connaissance des Catopidae (Coleoptera) paléarctiques. Acta Zoologica Cracoviensia, 15, 259–282.
Tinaut, A. (1998) Artrópodos terrestres de las cavidades andaluzas. Zoologica Baetica, 9, 3–28.
Tizado, E.J. & Salgado, J.M. (2000) Local-scale distribution of cholevid beetles (Col., Leiodidae: Cholevinae) in the province of León. Acta Oecologica, 21, 29–35.
https://doi.org/10.1016/S1146-609X(00)00109-0
Tizado, E.J., Salgado, J.M. & Núñez, E. (1995) Ecological correlates of the distribution of species of the family Cholevidae (Coleoptera) in the Cantabrian Mountains of northwest Spain. Acta Oecologica, 16, 351–360.
Torres, T., Ortiz, J.E., Soler, V., Reyes, E., Delgado, A., Valle, M., Llamas, J.F., Cobo, R., Julià, R., Badal, E., García de la Morena, M.A., García-Martínez, M.J., Fernández-Gianotti, J., Calvo, J.P. & Cortés, A. (2003) Pleistocene lacustrine basin of the east domain of guadix-baza basin (Granada, Spain): sedimentology, chronostratigraphy, and palaeoenvironment. In: Valero-Garcés, B.L. (Ed.), Limnogeología en España: un homenaje a Kerry R. Kelts. Consejo Superior de Investigaciones Científicas, Madrid, pp. 151–185.
Uhagón, S. (1890) Ensayo sobre las especies españolas del grupo "Cholevae". Anales de la Sociedad Española de Historia Natural, 19, 15–96.
Uhagón, S. (1898) Adiciones a mi "Ensayo sobre las especies españolas del grupo Cholevae". Anales de la Sociedad Española de Historia Natural, 7 (2), 27 (Actas de la Sociedad Española de Historia Natural, sesión del 1º de VI-1898), 117–126.
Velo-Antón, G., Godinho, R., Harris, D.J., Santos, X., Martínez-Freiria, F., Fahd, S., Larbes, S., Pleguezuelos, J.M. & Brito, J.C. (2012) Deep evolutionary lineages in a Western Mediterranean snake (Vipera latastei/monticola group) and high genetic structuring in Southern Iberian populations. Molecular Phylogenetics and Evolution, 65, 965–973.
https://doi.org/10.1016/j.ympev.2012.08.016
Vogler, A.P., DeSalle, R., Assmann, T., Knisley, C.B. & Schultz, T.D. (1993) Molecular population genetics of the endangered tiger beetle Cicindela dorsalis (Coleoptera, Cicindelidae). Annals of the Entomological Society of America, 86, 142–152.
https://doi.org/10.1093/aesa/86.2.142
Zangari, F., Cimmaruta, R. & Nascetti, G. (2006) Genetic relationships of the western Mediterranean painted frogs based on allozymes and mitochondrial markers: evolutionary and taxonomic inferences (Amphibia, Anura, Discoglossidae). Biological Journal of the Linnean Society, 87, 515–536.
https://doi.org/10.1111/j.1095-8312.2006.00585.x