Abstract
Seven new species of the genus Chrysometa Simon are described: C. citlaltepetl n. sp., C. triangulosa n. sp., C. rosarium n. sp., C. atotonilco n. sp., C. xamaticpac n. sp. C. puya n. sp. and C. sagicuta n. sp. Species identities were evaluated and sexes for each species matched with a fragment of the cytochrome c oxidase subunit I. These data were analyzed with maximum likelihood and the resulting cladograms separated all species with high support values (95–100) and an average distance of 0.093 %. The genetic signal also agreed with the diagnostic morphological features used to separate these taxa. The sex matching results discovered that the female of C. chipinque Levi actually belongs to C. puya n. sp.; the correct female of C. chipinque is here described for the first time. A redescription of the male of C. chipinque and the female of C. puya is also provided. All species were collected as part of a faunistic inventory from two oak forests near Pico de Orizaba Volcano National Park. A total of 399 adult specimens, 209 females and 195 males, were sorted and identified. Most individuals were collected from medium height vegetation by beating trays and from high vegetation by direct collecting at night. High resolution images for all species are available at www.unamfcaracnolab.com. Finally, the functional anatomy of the epigynum for the species described here is discussed.
References
Agnarsson, I., Gregoric, M., Blackledge, T.A. & Kuntner, M. (2012) The phylogenetic placement of Psechridae within Entelegynae and the convergent origin of orb-like spider webs.Journal of Zoological Systematics and Evolutionary Research, 51, 100–106.
https://doi.org/10.1111/jzs.12007Álvarez-Padilla Laboratory (2017) Araneomorphae of Mexico, a Digital Images Catalog. Facultad de Ciencias UNAM. Available from: http://www.unamfcaracnolab.com (accessed 1 May 2017)
Álvarez-Padilla, F. (2007) Systematics of the spider genus Metabus O.P-Cambridge, 1899 (Araneoidea: Tetragnathidae) with additions to the tetragnathid fauna of Chile and comments of the phylogeny of Tetragnathidae. Zoological Journal of the Linnean Society, 151, 285–335.
https://doi.org/10.1111/j.1096-3642.2007.00304.xÁlvarez-Padilla, F. & Hormiga, G. (2007) A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy. Journal of Arachnology, 35, 538–542.
https://doi.org/10.1636/Sh06-55.1Álvarez-Padilla, F., Dimitrov, D., Giribet, G. & Hormiga, G. (2009) Phylogenetic relationships of the spider family Tetragnathidae (Araneae, Araneoidea) based on morphological and DNA sequence data. Cladistics, 25, 109–146.
https://doi.org/10.1111/j.1096-0031.2008.00242.xÁlvarez-Padilla, F. & Hormiga, G. (2011) Morphological and phylogenetic atlas of the orb-weaving spider family Tetragnathidae (Araneae: Araneoidea). Zoological Journal of the Linnean Society, 162, 713–879.
https://doi.org/10.1111/j.1096-3642.2011.00692.xAstrin, J.J., Huber, B.A., Misof, B. & Klütsch, C.F.C. (2006) Molecular taxonomy in pholcid spiders (Pholcidae, Araneae): Evaluation of species identification methods using CO1 and 16S rRNA. Zoologica Scripta, 35, 441–457.
https://doi.org/10.1111/j.1463-6409.2006.00239.xBarnes, R.D. (1953) The ecological distribution of spiders in non-forest maritime communities at Beaufort, North Carolina. Ecological Monographs, 23, 315–337.
https://doi.org/10.1111/j.1463-6409.2006.00239.xBarrett, R.D.H. & Hebert, P.D.N. (2005) Identifying spiders through DNA barcodes. Canadian Journal of Zoology, 83, 481–491.
https://doi.org/10.1139/z05-024Bayer, S. & Schönhofer, A.L. (2013) Phylogenetic relationships of the spider family Psechridae inferred from molecular data, with comments on the Lycosoidea (Arachnida: Araneae). Invertebrate Systematics, 27, 53–80.
https://doi.org/10.1071/IS12017Blackledge, T.A., Scharff, N., Coddington, J.A., Szüts, T., Wenzel, J.W., Hayashi, C.Y. & Agnarsson, I. (2009) Reconstructing web evolution and spider diversification in the molecular era. Proceedings of the National Academy of Sciences, USA, 106, 5229–5234.
https://doi.org/10.1073/pnas.0901377106Blagoev, G., Hebert, P., Adamowicz, S. & Robinson, E. (2009) Prospects for using DNA barcoding to identify spiders in species-rich genera. ZooKeys, 16, 27–46.
https://doi.org/10.3897/zookeys.16.239Blagoev, G.A., DeWaard, J.R., Ratnasingham, S., DeWaard, S.L., Lu, L., Robertson, J., Telfer, A.C. & Hebert, P.D.N. (2016) Untangling taxonomy: A DNA barcode reference library for Canadian spiders. Molecular Ecology Resources, 16, 325–341.
https://doi.org/10.1111/1755-0998.12444Blagoev, G.A., Nikolova, N.I., Sobel, C.N., Hebert, P.D.N. & Adamowicz, S.J. (2013) Spiders (Araneae) of Churchill, Manitoba: DNA barcodes and morphology reveal high species diversity and new Canadian records. BMC ecology, 13, 44.
https://doi.org/10.1186/1472-6785-13-44Cardoso, P., Gaspar, C., Pereira, L.C., Silva, I., Henriques, S.S., da Silva, R.R. & Sousa, P. (2008b) Assessing spider species richness and composition in Mediterranean cork oak forests. Acta Oecologica, 33, 114–127.
https://doi.org/10.1016/j.actao.2007.10.003Cardoso, P., Scharff, N., Gaspar, C., Henriques, S.S., Carvalho, R., Castro, P.H., Schmidt, J.B., Silva, I., Szüts, T., de Castro, A. & Crespo, L.C. (2008a) Rapid biodiversity assessment of spiders (Araneae) using semi-quantitative sampling: a case study in a Mediterranean forest. Insect Conservation and Diversity, 1, 71–84.
https://doi.org/10.1111/j.1752-4598.2007.00008.xCastalanelli, M.A., Teale, R., Rix, M.G., Kennington, W.J. & Harvey, M.S. (2014) Barcoding of mygalomorph spiders (Araneae: Mygalomorphae) in the Pilbara bioregion of Western Australia reveals a highly diverse biota. Invertebrate Systematics, 28, 375–385.
Coddington, J. (1983) A temporary slide mount allowing precise manipulation of small structures. Verhandlungen des naturwissenschaftlichen Vereins Hamburg, 26, 291–292.
Coddington, J.A., Griswold, C.E., Silva-Dávila, D., Peñaranda, E. & Larcher, S.F. (1991) Designing and testing sampling protocols to estimate biodiversity in tropical ecosystems. In: Dudley, E.C. (Ed.), The Unity of Evolutionary Biology. Proceedings of the Fourth International Congress of Systematic and Evolutionary Biology, 1991, pp. 44–60.
CONANP (2015) Programa de manejo Parque Nacional El Pico de Orizaba. Comisión Nacional de Áreas Naturales Protegidas, 2015, 1–192. [in Spanish]
Cummings, M.P., Neel, M.C. & Shaw, K.L. (2008) A genealogical approach to quantifying lineage divergence. Evolution, 62, 2411–2422.
https://doi.org/10.1111/j.1558-5646.2008.00442.xDimitrov, D. & Hormiga, G. (2011) An extraordinary new genus of spiders from Western Australia with an expanded hypothesis on the phylogeny of Tetragnathidae (Araneae). Zoological Journal of the Linnean Society, 161, 735–768.
https://doi.org/10.1111/j.1096-3642.2010.00662.xDimitrov, D., Astrin, J.J. & Huber, B.A. (2013) Pholcid spider molecular systematics revisited, with new insights into the biogeography and the evolution of the group. Cladistics, 29, 132–146.
https://doi.org/10.1111/j.1096-0031.2012.00419.xDimitrov, D., Benavides, L.R., Arnedo, M.A., Giribet, G., Griswold, C.E., Scharff, N. & Hormiga, G. (2017) Rounding up the usual suspects: a standard target-gene approach for resolving the interfamilial phylogenetic relationships of ecribellate orb-weaving spiders with a new family-rank classification (Araneae, Araneoidea). Cladistics, 33, 221–250.
https://doi.org/10.1111/cla.12165Dimitrov, D., Lopardo, L., Giribet, G., Arnedo, M.A., Alvarez-Padilla, F. & Hormiga, G. (2012) Tangled in a sparse spider web: single origin of orb weavers and their spinning work unraveled by denser taxonomic sampling.Proceedings of the Royal Society of London B, 279, 1341–1350.
https://doi.org/10.1098/rspb.2011.2011Espuny, A., Barrientos, J.A. & Ascaso, C. (1993) Arañas de un encinar montano (Montseny, Barcelona, España). Resultados faunísticos. Bollettino Accademia Gioenia Scienze Naturali, 26, 93–105.
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.
González, R. (1993) La diversidad de los encinos mexicanos. Revista de la Sociedad Mexicana de Historia Natural, 44, 125–142.
Grandjean, F. (1949) Observation et conservation des très petits arthropodes. Bulletin du Muséum National d’Histoire naturelle, 21, 363–370.
Hamilton, C.A., Hendrixson, B.E., Brewer, M.S. & Bond, J.E. (2014) An evaluation of sampling effects on multiple DNA barcoding methods leads to an integrative approach for delimiting species: A case study of the North American tarantula genus Aphonopelma (Araneae, Mygalomorphae, Theraphosidae). Molecular Phylogenetics and Evolution, 71, 79–93.
https://doi.org/10.1016/j.ympev.2013.11.007Hebert, P.D.N., Cywinska, A., Ball, S.L. & deWaard, J.R. (2003a) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London. Series B, Biological Sciences, 270, 313–321.
https://doi.org/10.1098/rspb.2002.2218Ivanova, N.V., deWaard, J.R. & Hebert, P.D.N. (2006) An inexpensive, automation-friendly protocol for recovering high-quality DNA. Molecular Ecology Resources, 6, 998–1002.
https://doi.org/10.1111/j.1471-8286.2006.01428.xKallal, R.J. & Hormiga, G. (2016) Redescription of the Jamaican orb-weaving spider Chrysometa pecki Levi, 1986 (Araneae, Tetragnathidae), including the first description of the male. Zootaxa, 4066 (1), 95–100.
https://doi.org/10.11646/zootaxa.4066.1.11Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C., Thierer, T., Ashton, B., Mentjies, P. & Drummond, A. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.
https://doi.org/10.1093/bioinformatics/bts199Koponen, S. (1996) Spiders (Araneae) on trunks and large branches of oak (Quercus robur) in SW Finland. Revue Suisse de Zoologie, Hors Série, 335–340.
Levi, H. (1986) The Neotropical orb-weaver genera Chrysometa and Homalometa (Araneae: Tetragnathidae). Bulletin of the Museum of Comparative Zoology, 151, 91–215.
Lopardo, L., Giribet, G. & Hormiga, G. (2011) Morphology to the rescue: molecular data and the signal of morphological characters in combined phylogenetic analyses–a case study from mysmenid spiders (Araneae, Mysmenidae), with comments on the evolution of web architecture. Cladistics, 27, 278–330.
https://doi.org/10.1111/j.1096-0031.2010.00332.xLopardo, L. & Hormiga, G. (2015) Out of the twilight zone: phylogeny and evolutionary morphology of the orb-weaving spider family Mysmenidae, with a focus on spinneret spigot morphology in symphytognathoids (Araneae, Araneoidea).Zoological Journal of the Linnean Society, 173, 527–786.
https://doi.org/10.1111/zoj.12199Lopardo, L. & Uhl, G. (2014) Testing mitochondrial marker efficacy for DNA barcoding in spiders: A test case using the dwarf spider genus Oedothorax (Araneae: Linyphiidae: Erigoninae). Invertebrate Systematics, 28, 501–521.
https://doi.org/10.1071/IS14017Masters, B. C., Fan, V. & Ross, H. A. (2011) Species Delimitation–a geneious plugin for the exploration of species boundaries. Molecular Ecology Resources, 11, 154–157.
https://doi.org/10.1111/j.1755-0998.2010.02896.xMichalak, S. (2012) RaxmlGUI version 1.5: a graphical front-end for RAxML. Organisms Diversity and Evolution, 12, 335–337.
https://doi.org/10.1007/s13127-011-0056-0Miller, J.A., Carmichael, A., Ramírez, M.J., Spagna, J.C., Haddad, C.R., Rezac, M., Johannesen, J., Kral, J., Wang, X. & Griswold, C.E. (2010) Phylogeny of entelegyne spiders: affinities of the family Penestomidae (new rank), generic phylogeny of Eresidae, and asymmetric rates of change in spinning organ evolution (Araneae, Araneoidea, Entelegynae).Molecular Phylogenetics and Evolution, 55, 786–804.
https://doi.org/10.1016/j.ympev.2010.02.021Miller, J.A., Griswold, C.E., Scharff, N., Rezac, M., Szuts, T. & Marharbai, M. (2012) The velvet spiders: an atlas of the Eresidae (Arachnida, Araneae). ZooKeys, 195, 1–144.
https://doi.org/10.3897/zookeys.195.2342Moradmand, M., Schönhofer, A.L. & Jäger, P. (2014) Molecular phylogeny of the spider family Sparassidae with focus on the genus Eusparassus and notes on the RTA-clade and ‘Laterigradae’.Molecular Phylogenetics and Evolution, 74, 48–65.
https://doi.org/10.1016/j.ympev.2014.01.021Myers, N., Mittermeier, R.A., Mittermeier, C.G., da Fonseca G.A.B. & Kent, J. (2000) Biodiversity hotspots for conservation priorities. Nature, 403, 853–858.
https://doi.org/10.1038/35002501Nixon, K.C. (2006) Global and Neotropical distribution and diversity of oak (genus Quercus) and oak forests ecological studies. In: Kappelle, M. (Ed.), Ecology and Conservation of Neotropical Montane Oak Forests. Springer, Berlin, pp. 3–13.
https://doi.org/10.1007/3-540-28909-7_1Nogueira, A.A., Pena-Barbosa, J.P.P., Venticinque, E.M. & Brescovit, A.D. (2011) The spider genus Chrysometa (Araneae, Tetragnathidae) from the Pico da Neblina and Serra do Tapirapecó mountains (Amazonas, Brazil): new species, new records, diversity and distribution along two altitudinal gradients. Zootaxa, 2772, 33–51.
Polotow, D., Carmichael, A. & Griswold, C.E. (2015) Total evidence analysis of the phylogenetic relationships of Lycosoidea spiders (Araneae, Entelegynae).Invertebrate Systematics, 29, 124–163.
https://doi.org/10.1071/IS14041Pons, 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, 595609.
https://doi.org/10.1080/10635150600852011Rix, M.G., Harvey, M. & Roberts, J. (2008) Molecular phylogenetics of the spider family Micropholcommatidae (Araneae: Araneae) using nuclear rRNA genes (18S and 28S).Molecular Phylogenetics and Evolution, 46, 1031–1048.
https://doi.org/10.1016/j.ympev.2007.11.001Rzedowski, J. (1978) La vegetación de México. Limusa, Mexico City, 432 pp.
Scharff, N., Coddington, J.A., Griswold, C.E., Hormiga, G. & Bjørn, P. de P. (2003) When to quit? Estimating spider species richness in a Northern European deciduous forest. The journal of Arachnology, 31, 246–273.
https://doi.org/10.1636/0161-8202(2003)031[0246:WTQESS]2.0.CO;2Simó, M., Álvarez, L. & Laborda, Á. (2016) The orb-weaving spider genus Chrysometa in Uruguay: distribution and description of a new species (Araneae, Tetragnathidae). Zootaxa, 4067 (5), 589–593.
https://doi.org/10.11646/zootaxa.4067.5.7Simon, 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/87.6.651Stamatakis, A., Hoover, P. & Rougemont, J. (2008) A Rapid Bootstrap Algorithm for the RAxML Web Servers. Systematics Biology, 57, 758–771.
https://doi.org/10.1080/10635150802429642Urones, C., Jerardino, M. & Fernández, J.L. (1990) Estudio ecológico de las arañas epigeas (Araneae) en un encinar adehesado de Quercus ilex subsp. ballota (Desf.) Samp. (provincia de Salamanca, España). Boletín de la Asociación Española de Entomología, 14, 185–197.
Valencia, S. (2004) Diversidad del género Quercus (Fagaceae) en México. Boletín de la Sociedad Botánica de México, 75, 33–53.
Wheeler, W.C., Coddington, J.A., Crowley, L.M., Dimitrov, D., Goloboff, P.A., Griswold, C.E., Hormiga, G., Prendini, L., Ramírez, M.J., Sierwald, P., Almeida-Silva, L., Alvarez-Padilla, F., Arnedo, M. A., Benavides Silva, L.R., Benjamin, S.P., Bond, J.E., Grismado, C.J., Hasan, E., Hedin, M., Izquierdo, M.A., Labarque, F.M., Ledford, J., Lopardo, L., Maddison, W.P., Miller, J.A., Piacentini, L.N., Platnick, N.I., Polotow, D., Silva-Dávila, D., Scharff, N., Szüts, T., Ubick, D., Vink, C.J., Wood, H.M. & Zhang, J. (2017) The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling. Cladistics, 33, 574–616.
https://doi.org/10.1111/cla.12182Wood, H.M., Griswold, C.E. & Gillespie, R. (2012) Phylogenetic placement of pelican spiders (Archaeidae, Araneae), with insight into evolution of the “neck” and predatory behaviors of the superfamily Palpimanoidea. Cladistics, 28, 598–626.
https://doi.org/10.1111/j.1096-0031.2012.00411.xWorld Spider Catalog (2017) World Spider Catalog. Version 18. Natural History Museum Bern. Available from: http://wsc.nmbe.ch (accessed 1 May 2017)