Abstract
The integration of independent data sets could solve problems in both traditional and DNA-based taxonomy. The aim of this study is to investigate the power of CO1 sequences and of morphometrics to distinguish closely related species in the spider genus Araniella. We put special emphasis on the species pair A. cucurbitina (Clerck, 1757) and A. opisthographa (Kulczyński, 1905) since the females are morphologically difficult to distinguish and often misidentified. A total of 216 sequences of eight Araniella species from seven European countries, North America and Asia were included in the molecular analysis. The results from both maximum likelihood and Bayesian phylogenetic inference indicate successful separation of six out of eight Araniella species, including A. cucurbitina and A. opisthographa. For the same six species, we detect no overlap of intra- and interspecific genetic divergence, leading to successful species identification with a threshold approach. In addition, morphometric analysis of the epigyna of A. cucurbitina and A. opisthographa supports species separation by two best explanatory ratios: receptaculum length and distance between receptaculum and copulatory duct. Although a small overlap in the ratios exists, the species identification rate increases when combining morphometric and molecular data, which demonstrates the efficiency of integrative approaches for distinguishing closely related species. However, none of the molecular approaches was able to separate closely related A. alpica (L. Koch, 1869) and A. inconspicua (Simon, 1874) due to shared CO1 haplotypes. Considering the clear morphological separation of the males and different habitat preferences, incomplete lineage sorting or introgressive hybridization could have led to identical CO1 sequences. Therefore, DNA-barcoding must be thoroughly tested even within small homogenous genera of spiders.
References
Almquist, S. (2005) Swedish Araneae, part 1: families Atypidae to Hahniidae (Linyphiidae excluded). Insect Systematics & Evolution, Supplement, 62, 1–284.
Ballard, J.W.O. & Whitlock, M.C. (2004) The incomplete natural history of mitochondria. Molecular Ecology, 13 (4), 729–744.
http://dx.doi.org/10.1046/j.1365-294X.2003.02063.xBandelt, H.J., Forster, P. & Röhl, A. (1999) Median-joining networks for inferring intraspecific phylogenies. Molecular Biology and Evolution, 16 (1), 37–48.
http://dx.doi.org/10.1093/oxfordjournals.molbev.a026036Barrett, R.D. & Hebert, P.D. (2005) Identifying spiders through DNA barcodes. Canadian Journal of Zoology, 83 (3), 481–491.
Baur, H., Kranz-Baltensperger, Y., Cruaud, A., Rasplus, J.Y., Timokhov, A.V. & Gokhman, V.E. (2014) Morphometric analysis and taxonomic revision of Anisopteromalus Ruschka (Hymenoptera: Chalcidoidea: Pteromalidae)–an integrative approach. Systematic Entomology, 39 (4), 691–709.
http://dx.doi.org/10.1111/syen.12081Baur, H. & Leuenberger, C. (2011) Analysis of ratios in multivariate morphometry. Systematic Biology, 60, 813–825.
http://dx.doi.org/10.1093/sysbio/syr061Bayer, 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 (1), 53–80.
http://dx.doi.org/10.1071/IS12017Bergsten, J., Bilton, D.T., Fujisawa, T., Elliott, M., Monaghan, M.T., Balke, M., Hendrich, L., Geijer, J., Herrmann, J., Foster, G.N., Ribera, I., Nilsson, A.N., Barraclough, T.G. & Vogler, A.P. (2012) The effect of geographical scale of sampling on DNA barcoding. Systematic biology, 61 (5), 851–869.
http://dx.doi.org/10.1093/sysbio/sys037Berthier, P., Excoffier, L. & Ruedi, M. (2006) Recurrent replacement of mtDNA and cryptic hybridization between two sibling bat species Myotis myotis and Myotis blythii. Proceedings of the Royal Society B: Biological Sciences, 273 (1605), 3101–3123.
http://dx.doi.org/10.1098/rspb.2006.3680Blagoev, G., Hebert, P., Adamowicz, S. & Robinson, E. (2009) Prospects for using DNA barcoding to identify spiders in species-rich genera. ZooKeys, 16, 27–46.
http://dx.doi.org/10.3897/zookeys.16.239Blagoev, G.A. & Dondale, C.D. (2014) A new species of Alopecosa (Araneae: Lycosidae) from Canada: a morphological description supported by DNA barcoding of 19 congeners. Zootaxa, 3894 (1), 152–160.
http://dx.doi.org/10.11646/zootaxa.3894.1.12Blagoev, G.A., Nikolova, N.I., Sobel, C.N., Hebert, P.D. & Adamowicz, S.J. (2013) Spiders (Araneae) of Churchill, Manitoba: DNA barcodes and morphology reveal high species diversity and new Canadian records. BMC Ecology, 13 (1), 1.
http://dx.doi.org/10.1186/1472-6785-13-44Blanke, R. (1982) Untersuchungen zur Taxonomie der Gattung Araniella (Araneae, Araneidae). Zoologica Scripta, 11 (4), 287–305.
http://dx.doi.org/10.1111/j.1463-6409.1982.tb00540.xBoyer, S.L., Baker, J.M. & Giribet, G. (2007) Deep genetic divergences in Aoraki denticulata (Arachnida, Opiliones, Cyphophthalmi): a widespread ‘mite harvestman’defies DNA taxonomy. Molecular Ecology, 16 (23), 4999–5016.
http://dx.doi.org/10.1111/j.1365-294X.2007.03555.xBrower, A.V. (2006) Problems with DNA barcodes for species delimitation: ‘ten species’ of Astraptes fulgerator reassessed (Lepidoptera: Hesperiidae). Systematics and Biodiversity, 4 (2), 127–132.
http://dx.doi.org/10.1017/S147720000500191XBrown, S.D. & Collins, R.A. (2011) Spider: Species Identity and Evolution in R. A Tutorial. Available from: http://spider.r-forge.r-project.org/tutorial/tutorial.pdf (accessed 1 May 2016)
Brown, S.D., Collins, R.A., Boyer, S., Lefort, M.C., Malumbres-Olarte, J., Vink, C.J. & Cruickshank, R.H. (2012) Spider: an R package for the analysis of species identity and evolution, with particular reference to DNA barcoding. Molecular Ecology Resources, 12 (3), 562–565.
http://dx.doi.org/10.1111/j.1755-0998.2011.03108.xČandek, K. & Kuntner, M. (2015) DNA barcoding gap: reliable species identification over morphological and geographical scales. Molecular Ecology Resources, 15 (2), 268–277.
http://dx.doi.org/10.1111/1755-0998.12304Cardoso, A. & Vogler, A. (2005) DNA taxonomy, phylogeny and Pleistocene diversification of the Cicindela hybrida species group (Coleoptera: Cicindelidae). Molecular Ecology, 14 (11), 3531–3546.
http://dx.doi.org/10.1111/j.1365-294X.2005.02679.xChamberlin, R.V. & Ivie, W. (1942) A hundred new species of American spiders. University of Utah, Utah, 117 pp.
Civetta, A., Pramual, P., Wongpakam, K. & Adler, P.H. (2010) Cryptic biodiversity and phylogenetic relationships revealed by DNA barcoding of Oriental black flies in the subgenus Gomphostilbia (Diptera: Simuliidae). Genome, 54 (1), 1–9.
Clerck, C. (1757) Svenska spindlar, uti sina hufvud-slågter indelte samt under några och sextio särskildte arter beskrefne och med illuminerade figurer uplyste. Literis Laur. Salvii, Stockholmiae, 154 pp.
Coddington, J.A., Agnarsson, I., Cheng, R., Čandek, K., Driskell, A., Frick, H., Gregorič, M., Kostanjšek, R., Kropf, C., Kweskin, M., Lokovšek, T., Pipan, M., Vidergar, N. & Kuntner, M. (2016) DNA barcode data accurately identify higher taxa. PeerJ PrePrints, 4, e1633v1.
https://doi.org/10.7287/peerj.preprints.1633v1Collins, R.A., Boykin, L.M., Cruickshank, R.H. & Armstrong, K.F. (2012) Barcoding's next top model: an evaluation of nucleotide substitution models for specimen identification. Methods in Ecology and Evolution, 3 (3), 457–465.
http://dx.doi.org/10.1111/j.2041-210X.2011.00176.xCooperation, C. (2011) North American Environmental Atlas. Available from: http://www.cec.org/naatlas (accessed 23 June 2015)
Croucher, P.J.P., Oxford, G.S. & Searle, J.B. (2004) Mitochondrial differentiation, introgression and phylogeny of species in the Tegenaria atrica group (Araneae: Agelenidae). Biological Journal of the Linnean Society, 81 (1), 79–89.
http://dx.doi.org/10.1111/j.1095-8312.2004.00280.xDayrat, B. (2005) Towards integrative taxonomy. Biological Journal of the Linnean Society, 85 (3), 407–415.
http://dx.doi.org/10.1111/j.1095-8312.2005.00503.xde Mendonça, R.S., Navia, D., Diniz, I.R., Auger, P. & Navajas, M. (2011) A critical review on some closely related species of Tetranychus sensu stricto (Acari: Tetranychidae) in the public DNA sequences databases. Experimental and Applied Acarology, 55 (1), 1–23.
http://dx.doi.org/10.1007/s10493-011-9453-5DeSalle, R., Egan, M.G. & Siddall, M. (2005) The unholy trinity: taxonomy, species delimitation and DNA barcoding. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 360 (1462), 1905–1916.
http://dx.doi.org/10.1098/rstb.2005.1722Elias, M., Hill, R.I., Willmott, K.R., Dasmahapatra, K.K., Brower, A.V., Mallet, J. & Jiggins, C.D. (2007) Limited performance of DNA barcoding in a diverse community of tropical butterflies. Proceedings of the Royal Society B: Biological Sciences, 274 (1627), 2881–2889.
http://dx.doi.org/10.1098/rspb.2007.1035Ferri, E., Barbuto, M., Bain, O., Galimberti, A., Uni, S., Guerrero, R., Ferté, H., Bandi, C., Martin, C. & Casiraghi, M. (2009) Integrated taxonomy: traditional approach and DNA barcoding for the identification of filarioid worms and related parasites (Nematoda). Frontiers in Zoology, 6 (1).
http://dx.doi.org/10.1186/1742-9994-6-1
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 (5), 294–299.
Fonseca, G., Derycke, S. & Moens, T. (2008) Integrative taxonomy in two free‐living nematode species complexes. Biological Journal of the Linnean Society, 94 (4), 737–753.
http://dx.doi.org/10.1111/j.1095-8312.2008.01015.xFontaneto, D., Kaya, M., Herniou, E.A. & Barraclough, T.G. (2009) Extreme levels of hidden diversity in microscopic animals (Rotifera) revealed by DNA taxonomy. Molecular Phylogenetics and Evolution, 53 (1), 182–189.
http://dx.doi.org/10.1016/j.ympev.2009.04.011Fregin, S., Haase, M., Olsson, U. & Alström, P. (2012) Pitfalls in comparisons of genetic distances: a case study of the avian family Acrocephalidae. Molecular Phylogenetics and Evolution, 62 (1), 319–328.
http://dx.doi.org/10.1016/j.ympev.2011.10.003Godfray, H.C.J. (2002) Challenges for taxonomy. Nature, 417 (6884), 17–19.
http://dx.doi.org/10.1038/417017aGoto, S.G. & Kimura, M.T. (2001) Phylogenetic utility of mitochondrial COI and nuclear Gpdh genes in Drosophila. Molecular Phylogenetics and Evolution, 18 (3), 404–422.
http://dx.doi.org/10.1006/mpev.2000.0893Greenstone, M., Rowley, D., Heimbach, U., Lundgren, J., Pfannenstiel, R. & Rehner, S. (2005) Barcoding generalist predators by polymerase chain reaction: carabids and spiders. Molecular Ecology, 14 (10), 3247–3266.
http://dx.doi.org/10.1111/j.1365-294X.2005.02628.xGregoric, M., Blackledge, T.A., Agnarsson, I. & Kuntner, M. (2015) A molecular phylogeny of bark spiders reveals new species from Africa and Madagascar (Araneae: Araneidae: Caerostris). Journal of Arachnology, 43 (3), 293–312.
http://dx.doi.org/10.1636/0161-8202-43.3.293Guindon, 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 (3), 307–321.
http://dx.doi.org/10.1093/sysbio/syq010Hajibabaei, M., Singer, G.A., Hebert, P.D. & Hickey, D.A. (2007) DNA barcoding: how it complements taxonomy, molecular phylogenetics and population genetics. Trends in Genetics, 23 (4), 167–172.
http://dx.doi.org/10.1016/j.tig.2007.02.001Hazewinkel, M. (2001) Student test. Encyclopedia of Mathematics. Springer, Berlin, 978 pp.
Hebert, P.D., Cywinska, A. & Ball, S. L. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society of London B: Biological Sciences, 270, 313–321.
http://dx.doi.org/10.1098/rspb.2002.2218Hebert, P.D. & Gregory, T.R. (2005) The promise of DNA barcoding for taxonomy. Systematic Biology, 54, 852–859.
http://dx.doi.org/10.1080/10635150500354886Hebert, P.D., Stoeckle, M.Y., Zemlak, T.S. & Francis, C.M. (2004) Identification of birds through DNA barcodes. PLoS Biology, 2, e312.
http://dx.doi.org/10.1371/journal.pbio.0020312Hedin, M.C. & Maddison, W.P. (2001) A combined molecular approach to phylogeny of the jumping spider subfamily Dendryphantinae (Araneae: Salticidae). Molecular Phylogenetics and Evolution, 18, 386–403.
http://dx.doi.org/10.1006/mpev.2000.0883Heimer, S., Nentwig, W. & Bosmans, R. (1991) Spinnen Mitteleuropas: Ein Bestimmungsbuch. P. Parey, Berlin, 543 pp.
Heinrichs, J., Kreier, H.-P., Feldberg, K., Schmidt, A.R., Zhu, R.-L., Shaw, B., Shaw, A.J. & Wissemann, V. (2011) Formalizing morphologically cryptic biological entities: new insights from DNA taxonomy, hybridization, and biogeography in the leafy liverwort Porella platyphylla (Jungermanniopsida, Porellales). American Journal of Botany, 98, 1252–1262.
http://dx.doi.org/10.3732/ajb.1100115Huber, B.A. & Dimitrov, D. (2014) Slow genital and genetic but rapid non-genital and ecological differentiation in a pair of spider species (Araneae, Pholcidae). Zoologischer Anzeiger-A Journal of Comparative Zoology, 253, 394–403.
http://dx.doi.org/10.1016/j.jcz.2014.04.001Johnson, S.B., Warén, A. & Vrijenhoek, R.C. (2008) DNA barcoding of Lepetodrilus limpets reveals cryptic species. Journal of Shellfish Research, 27, 43–51.
http://dx.doi.org/10.2983/0730-8000(2008)27%5B43:DBOLLR%5D2.0.CO;2Kaila, L. & Ståhls, G. (2006) DNA barcodes: Evaluating the potential of COI to diffentiate closely related species of Elachista (Lepidoptera: Gelechioidea: Elachistidae) from Australia. Zootaxa, 1170, 1–26.
Kearse, M., Moir, R., Wilson, A., Stones-Havas, S., Cheung, M., Sturrock, S., Buxton, S., Cooper, A., Markowitz, S., Duran, C. & Thierer, T. (2012) Geneious Basic: an integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics, 28, 1647–1649.
http://dx.doi.org/10.1093/bioinformatics/bts199Koch, L. (1869) Beitrag zur Kenntniss der Arachnidenfauna Tirols. Zeitschrift des Ferdinandeums für Tirol und Vorarlberg, Series 3, 14, 149–206.
Kulczyński, W. (1885) Araneae in Camtschadalia a Dre B. Dybowski collectae. Pamietnik Akademji umiejetnosci w Krakow wydzial matematyczno-przyrodniczy, 11, 1–60.
Kulczyński, W. (1905) Fragmenta arachnologica. V. Bulletin International de l'Academie des Sciences de Cracovie, 231–250.
Kumar, N.P., Rajavel, A., Natarajan, R. & Jambulingam, P. (2007) DNA barcodes can distinguish species of Indian mosquitoes (Diptera: Culicidae). Journal of Medical Entomology, 44, 01–07.
http://dx.doi.org/10.1093/jmedent/41.5.01Laszlo, Z., Baur, H. & Tothmeresz, B. (2013) Multivariate ratio analysis reveals Trigonoderus pedicellaris Thomson (Hymenoptera, Chalcidoidea, Pteromalidae) as a valid species. Systematic Entomology, 38, 753–762.
http://dx.doi.org/10.1111/syen.12026Levi, H.W. (1974) The orb-weaver genera Araniella and Nuctenea (Araneae: Araneidae). Bulletin of the Museum of Comparative Zoology at Harvard College, 146, 291–316.
Levy, G. (1987) Spiders of the genera Araniella, Zygiella, Zilla and Mangora (Araneae, Araneidae) from Israel, with notes on Metellina species from Lebanon. Zoologica Scripta, 16, 243–257.
http://dx.doi.org/10.1111/j.1463-6409.1987.tb00071.xLipscomb, D., Platnick, N. & Wheeler, Q. (2003) The intellectual content of taxonomy: a comment on DNA taxonomy. Trends in Ecology & Evolution, 18 (2), 65–66.
http://dx.doi.org/10.1016/S0169-5347(02)00060-5Maddison, W.P. (1997) Gene trees in species trees. Systematic Biology, 46, 523–536.
http://dx.doi.org/10.1093/sysbio/46.3.523Mallet, J. (2001) Species, concepts of. Encyclopedia of biodiversity, 5, 427–440.
http://dx.doi.org/10.1016/B0-12-226865-2/00254-6Mallet, J. (2005) Hybridization as an invasion of the genome. Trends in Ecology & Evolution, 20, 229–237.
http://dx.doi.org/10.1016/j.tree.2005.02.010Mallet, J. (2008) Hybridization, ecological races and the nature of species: empirical evidence for the ease of speciation. Philosophical Transactions of the Royal Society of London B: Biological Sciences, 363, 2971–2986.
http://dx.doi.org/10.1098/rstb.2008.0081Mallet, J. & Willmott, K. (2003) Taxonomy: renaissance or Tower of Babel? Trends in Ecology & Evolution, 18, 57–59.
http://dx.doi.org/10.1016/S0169-5347(02)00061-7Meier, R., Shiyang, K., Vaidya, G. & Ng, P.K. (2006) DNA barcoding and taxonomy in Diptera: a tale of high intraspecific variability and low identification success. Systematic Biology, 55, 715–728.
http://dx.doi.org/10.1080/10635150600969864Meyer, C.P. & Paulay, G. (2005) DNA barcoding: error rates based on comprehensive sampling. PLoS Biology, 3, e422.
http://dx.doi.org/10.1371/journal.pbio.0030422Monaghan, M.T., Balke, M., Pons, J. & Vogler, A.P. (2006) Beyond barcodes: complex DNA taxonomy of a South Pacific Island radiation. Proceedings of the Royal Society B: Biological Sciences, 273, 887–893.
http://dx.doi.org/10.1098/rspb.2005.3391Moritz, C. & Cicero, C. (2004) DNA barcoding: promise and pitfalls. PLoS Biology, 2, e354.
http://dx.doi.org/10.1371/journal.pbio.0020354Mutanen, M., Kekkonen, M., Prosser, S.W., Hebert, P.D. & Kaila, L. (2015) One species in eight: DNA barcodes from type specimens resolve a taxonomic quagmire. Molecular Ecology Resources. 5 (4), 967–984.
http://dx.doi.org/10.1111/1755-0998.12361Nadolny, A.A., Omelko, M.M., Marusik, Y.M., & Blagoev, G. (2016) A new species of spider belonging to the Pardosa lugubris-group (Araneae: Lycosidae) from Far East Asia. Zootaxa, 4072 (2), 263–281.
http://dx.doi.org/10.11646/zootaxa.4072.2.8Namkung, J. (2002) The spiders of Korea. Kyo-Hak Publishing Co., Seoul, 648 pp.
Nentwig, W., Blick, T., Gloor, D., Hänggi, A. & Kropf, C. (2010) Spiders of Europe. Available from: http://www.araneae.unibe.ch (accessed 21 January 2015)
Packer, L., Gibbs, J., Sheffield, C. & Hanner, R. (2009) DNA barcoding and the mediocrity of morphology. Molecular Ecology Resources, 9, 42–50.
http://dx.doi.org/10.1111/j.1755-0998.2009.02631.xPadial, J.M., Miralles, A., De la Riva, I. & Vences, M. (2010) Review: The integrative future of taxonomy. Frontiers in Zoology, 7, 1–14.
http://dx.doi.org/10.1186/1742-9994-7-16Paquin, P. & Hedin, M. (2004) The power and perils of ‘molecular taxonomy’: a case study of eyeless and endangered Cicurina (Araneae: Dictynidae) from Texas caves. Molecular Ecology, 13 (10), 3239–3255.
http://dx.doi.org/10.1111/j.1365-294X.2004.02296.xPina, T., Verdú, M.J., Urbaneja, A. & Sabater-Muñoz, B. (2012) The use of integrative taxonomy in determining species limits in the convergent pupa coloration pattern of Aphytis species. Biological Control, 61, 64–70.
http://dx.doi.org/10.1016/j.biocontrol.2011.11.004Planas, E. & Ribera, C. (2015) Description of six new species of Loxosceles (Araneae: Sicariidae) endemic to the Canary Islands and the utility of DNA barcoding for their fast and accurate identification. Zoological Journal of the Linnean Society, 174 (1), 47–73.
http://dx.doi.org/10.1111/zoj.12226Prendini, L. (2005) Comment on "Identifying spiders through DNA barcodes". Canadian Journal of Zoology, 83 (3), 498–504.
http://dx.doi.org/10.1139/z05-025Quicke, D.L., Alex Smith, M., Janzen, D.H., Hallwachs, W., Fernandez-Triana, J., Laurenne, N.M., Zaldivar-Riveroni, A., Shaw, M.R., Broad, G.R., Klopfstein, S., Shaw, S.T., Hrcek, J., Hebert, P.D., Miller, S.E., Rodriguez, J.J., Whitfield, J.B., Sharkey, M.J, Sharanowski, B.J., Jussila, R., Gauld, I.D., Chesters, D. & Vogler, A.P. (2012) Utility of the DNA barcoding gene fragment for parasitic wasp phylogeny (Hymenoptera: Ichneumonoidea): data release and new measure of taxonomic congruence. Molecular Ecology Resources, 12, 676–685.
http://dx.doi.org/10.1111/j.1755-0998.2012.03143.xR Core Team (2013) R: A Language and Environment for Statistical Computing. Vienna, Austria, 2011. Avaiable from: http://www.R-project.orgRajaei Sh, H., Struwe, J.F., Raupach, M.J., Ahrens, D. & Wägele, J.W. (2013) Integration of cytochrome c oxidase I barcodes and geometric morphometrics to delimit species in the genus Gnopharmia (Lepidoptera: Geometridae, Ennominae). Zoological Journal of the Linnean Society, 169, 70–83.
Rambaut, A. (2009) FigTree v1. 3.1: Tree figure drawing tool. Avaiable from: http://tree.bio.ed.ac.uk/software/figtree (accessed 19 April 2016)
Rasban, W.S. (1997–2015) Image J V. 1.312 u, Image Processing and Analysis in Java. Avaiable from: http://imagej.nih.gov/ij/ (accessed 19 April 2016)
Ratnasingham, S. & Hebert, P.D. (2007) BOLD: The Barcode of Life Data System (http://www. barcodinglife. org). Molecular Ecology Notes, 7, 355–364.
http://dx.doi.org/10.1111/j.1471-8286.2007.01678.xŘezáč, M., Gasparo, F., Král, J. & Heneberg, P. (2014) Integrative taxonomy and evolutionary history of a newly revealed spider Dysdera ninnii complex (Araneae: Dysderidae). Zoological Journal of the Linnean Society, 172, 451–474.
Riedel, A., Sagata, K., Suhardjono, Y.R., Tänzler, R. & Balke, M. (2013) Integrative taxonomy on the fast track-towards more sustainability in biodiversity research. Frontiers in Zoology, 10, 15.
http://dx.doi.org/10.1186/1742-9994-10-15Roberts, M. & Jones, R. (1996) Collins field guide. Spiders of Britain and northern Europe. British Journal of Entomology and Natural History, 9, 195–195.
Roberts, M.J. (1987) The spiders of Great Britain and Ireland. Vol. 2. Brill Archive, Leiden, 204 pp.
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.
http://dx.doi.org/10.1093/sysbio/sys029Sacher, P. (1990) Untersuchungen über Zahl, Anordnung und taxonomischen Wert der dorso-lateralen Abdominalpunkte in der Gattung Araniella (Arachnida: Araneidae). Acta Zoologica Fennica, 190, 345–349.
Schlick-Steiner, B.C., Steiner, F.M., Seifert, B., Stauffer, C., Christian, E. & Crozier, R.H. (2010) Integrative taxonomy: a multisource approach to exploring biodiversity. Annual Review of Entomology, 55, 421–438.
http://dx.doi.org/10.1146/annurev-ento-112408-085432Schmidt, S., Schmid-Egger, C., Morinière, J., Haszprunar, G. & Hebert, P.D. (2015) DNA barcoding largely supports 250 years of classical taxonomy: identifications for Central European bees (Hymenoptera, Apoidea partim). Molecular Ecology Resources.
http://dx.doi.org/10.1111/1755-0998.12363Sheffield, C.S., Hebert, P.D., Kevan, P.G. & Packer, L. (2009) DNA barcoding a regional bee (Hymenoptera: Apoidea) fauna and its potential for ecological studies. Molecular Ecology Resources, 9, 196–207.
http://dx.doi.org/10.1111/j.1755-0998.2009.02645.xSimon, E. (1874) Les arachnides de France. Librairie Encyclopédique de Roret, Paris, 272 pp.
Srivathsan, A. & Meier, R. (2012) On the inappropriate use of Kimura-2-parameter (K2P) divergences in the DNA-barcoding literature. Cladistics, 28, 190–194.
http://dx.doi.org/10.1111/j.1096-0031.2011.00370.xStevens, P. (1991) Character states, morphological variation, and phylogenetic analysis: a review. Systematic Botany, 553–583.
http://dx.doi.org/10.2307/2419343Tanikawa, A. (1995) A revision of the Japanese spiders of the genus Araniella (Araneae: Araneidae). Acta Arachnologica, 44 (1), 51–60.
Tamura, K., Stecher, G., Peterson, D., Filipski, A. & Kumar, S. (2013) MEGA6: molecular evolutionary genetics analysis version 6.0. Molecular Biology and Evolution, 30, 2725–2729.
http://dx.doi.org/10.1093/molbev/mst197Tautz, D., Arctander, P., Minelli, A., Thomas, R.H. & Vogler, A.P. (2003) A plea for DNA taxonomy. Trends in Ecology & Evolution, 18, 70–74.
http://dx.doi.org/10.1016/S0169-5347(02)00041-1Teletchea, F. (2010) After 7 years and 1000 citations: comparative assessment of the DNA barcoding and the DNA taxonomy proposals for taxonomists and non-taxonomists. Mitochondrial DNA, 21, 206–226.
http://dx.doi.org/10.3109/19401736.2010.532212Trewick, S.A. (2008) DNA Barcoding is not enough: mismatch of taxonomy and genealogy in New Zealand grasshoppers (Orthoptera: Acrididae). Cladistics, 24, 240–254.
http://dx.doi.org/10.1111/j.1096-0031.2007.00174.xVan Der Bank, H., Herbert, D., Greenfield, R. & Yessoufou, K. (2013) Revisiting species delimitation within the genus Oxystele using DNA barcoding approach. ZooKeys, 337.
van Velzen, R., Weitschek, E., Felici, G. & Bakker, F.T. (2012) DNA barcoding of recently diverged species: relative performance of matching methods. PloS one, 7, e30490.
http://dx.doi.org/10.1371/journal.pone.0030490Wahlberg, N., Oliveira, R. & Scott, J.A. (2003) Phylogenetic relationships of Phyciodes butterfly species (Lepidoptera: Nymphalidae): complex mtDNA variation and species delimitations. Systematic Entomology, 28, 257–274.
http://dx.doi.org/10.1046/j.1365-3113.2003.00212.xWallman, J. & Donnellan, S. (2001) The utility of mitochondrial DNA sequences for the identification of forensically important blowflies (Diptera: Calliphoridae) in southeastern Australia. Forensic Science International, 120, 60–67.
http://dx.doi.org/10.1016/S0379-0738(01)00426-1Wheeler, Q.D. (2005) Losing the plot: DNA “barcodes” and taxonomy. Cladistics, 21 (4), 405–407.
http://dx.doi.org/10.1111/j.1096-0031.2005.00075.xWiehle, H. (1960) Die Tierwelt Deutschlands. 47. Spinnentiere oder Arachnoidea (Araneae), 11, 1–620.
Will, K.W., & Rubinoff, D. (2004) Myth of the molecule: DNA barcodes for species cannot replace morphology for identification and classification. Cladistics, 20 (1), 47–55.
http://dx.doi.org/10.1111/j.1096-0031.2003.00008.xWill, K.W., Mishler, B.D. & Wheeler, Q.D. (2005) The perils of DNA barcoding and the need for integrative taxonomy. Systematic Biology, 54, 844–851.
http://dx.doi.org/10.1080/10635150500354878World Spider Catalog (2016) World Spider Catalog. Natural History Museum Bern. Available from: http://wsc.nmbe.ch (accessed 8 February 2016)
Xia, X. (2013) DAMBE5: a comprehensive software package for data analysis in molecular biology and evolution. Molecular biology and evolution, 30, 1720–1728.
http://dx.doi.org/10.1093/molbev/mst064Xia, X. & Lemey, P. (2009) Assessing substitution saturation with DAMBE. The phylogenetic handbook: a practical approach to DNA and protein phylogeny, 2, 615–630.
http://dx.doi.org/10.1017/CBO9780511819049.022Xia, X., Xie, Z., Salemi, M., Chen, L. & Wang, Y. (2003) An index of substitution saturation and its application. Molecular Phylogenetics and Evolution, 26, 1–7.
http://dx.doi.org/10.1016/S1055-7903(02)00326-3