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Issue: Vol. 5624 No. 1: 23 Apr. 2025
Type: Monograph
Published: 2025-04-23
DOI: 10.11646/zootaxa.5624.1.1
Page range: 1-69
Abstract views: 15
PDF downloaded: 1

Integrative taxonomy of the Iberian Zodarion species of the rubidum and styliferum groups (Araneae: Zodariidae)

Department of Botany and Zoology, Faculty of Sciences, Masaryk University, Brno, Czech Republic
Department of Botany and Zoology, Faculty of Sciences, Masaryk University, Brno, Czech Republic
Department of Botany and Zoology, Faculty of Sciences, Masaryk University, Brno, Czech Republic
Araneae Ant-eating spiders COI copulatory organs description genetic divergence Mediterranean region phylogeny

Abstract

Zodarion, one of the most diverse zodariid spider genera, has recently garnered significant interest due to its peculiar adaptations for stringent ant-eating specialization. However, species identification within this genus remains challenging, as many Zodarion species are still insufficiently documented. This study revisits the taxonomy of Iberian species within the rubidum and styliferum groups, which together comprise 25 of the 171 species in the genus. Detailed morphological documentation is provided, including photographs and SEM images of key taxonomic characters, presented here for the first time. Molecular characterizations were also conducted to facilitate future species identification and to investigate patterns of mitochondrial variation within and between species. Notably, Z. extraneum has been reinstated as a distinct species. The male of Z. extraneum and the female of Z. alentejanum are described for the first time, while the female of Z. viduum is redescribed. Additionally, the species Z. alentejanum, Z. bosmansi, Z. guadianense and Z. lusitanicum have been assigned to the styliferum group. Three new synonyms are proposed: Z. brevicephalus with Z. styliferum, Z. duriense with Z. bacelarae, and Z. parastyliferum with Z. extraneum. Pairwise interspecific divergences between individual sequences generally exceeded intraspecific variation, and the mitochondrial gene tree recovered all but one species as monophyletic groups, underscoring the utility of molecular barcoding for identifying Zodarion species. However, interspecific distances between consensus sequences often blurred species boundaries. Our integrative taxonomic study provides tools for improved identification of Zodarion species, facilitating further research into their biology.

References

  1. Aberer, A.J., Krompass, D. & Stamatakis A. (2013) Pruning rogue taxa improves phylogenetic accuracy: an efficient algorithm and webservice. Systematic Biology, 62, 162–166. https://doi.org/10.1093/sysbio/sys078
  2. Askins, M. (1999) Zodarion fuscum (Simon, 1870): A spider new to Britain. Newsletter of the British Arachnological Society, 86, 11.
  3. Audouin, V. (1826) Explication sommaire des planches d’arachnides de l’Égypte et de la Syrie. In: Savigny, M.J.C.L. (Ed.), “Description de l’Égypte, ou recueil des observations et des recherches qui ont été faites en Égypte pendant l’expédition de l’armée franҫaise, publié par les ordres de sa Majesté l’Empereur Napoléon le Grand.”. Histoire Naturelle, 1 (4), 1–339.
  4. Bacelar, A. (1928) Aracnídios Portguêses: III. Bulletin da Sociedade portuguesa de Sciencias Naturais, 10, 169–203.
  5. Barrientos, J.A., Blasco, A., Ferrández, M.A., Godall, P., Pérez, J.A., Rambla, M. & Utones, C. (1985) Artrópodos epigeos del Macizo de San Juan de la Peña (Jaca, Huesca). XIII. Familias de Araneidos de escasa representacion. Pirineos, 35, 211–234.
  6. Barrientos, J.A., Moya, J., García-Sarrión, R., Uribarri, I., Melic, A., Prieto, C.E., Moraza, M.L. & Zaragoza, J.A. (2017) Arácnidos del Parque Natural de Cabo de Gata-Níjar (Almería, España). Revista Ibérica de Aracnología, 30, 107–138.
  7. Bosmans, R. (1994) Revision of the genus Zodarion Walckenaer, 1833 in the Iberian peninsula and Balearic islands (Araneae, Zodariidae). Eos, 69, 115–142.
  8. Bosmans, R. (1997) Revision of the genus Zodarion Walckenaer, 1833, part II. Western and Central Europe, including Italy (Araneae: Zodariidae). Bulletin-British Arachnological Society, 10 (8), 265–294.
  9. Bosmans, R. (2009) Revision of the genus Zodarion Walckenaer, 1833, part III. South East Europe and Turkey (Araneae: Zodariidae). Contributions to Natural History, 12, 1–211.
  10. Breitling, R. (2020) South European spiders from the Duffey collection in the Manchester Museum (Arachnida: Araneae). Arachnology, 18, 333–362. https://doi.org/10.13156/arac.2020.18.4.333
  11. Cárdenas, M., Jiroš, P. & Pekár, S. (2012) Selective olfactory attention of a specialised predator to intraspecific chemical signals of its prey. Naturwissenschaften, 99, 597–605. https://doi.org/10.1007/s00114-012-0938-9
  12. Crespo, L.C. (2008) Contribution to the knowledge of the Portuguese spider (Arachnida: Araneae) fauna: seven new additions to the Portuguese checklist. Boletín de la Sociedad Entomológica Aragonesa, 43, 403–407.
  13. Crespo, L.C., Cardoso, P., Henriques, S. & Gaspar, C. (2009) Spiders (Araneae) from Porto Santo (Madeira, Portugal): additions to the current knowledge. Boletín de la Sociedad Entomologica aragonesa, 45, 471–475.
  14. Denis, J. (1935a) A propos de quelques araignées du genre Zodarion Walck., appartenant à la faune française. Bulletin de la Société d’Histoire naturelle de Toulouse, 67, 51–68.
  15. Denis, J. (1935b) Sur deux araignées de Cyrénaique. Annali del Museo Civico di Storia Naturale di Genova, 57, 100–104.
  16. Denis, J. (1937) Contribution à l’étude des araignées du genre Zodarion Walckenaer. Festschrift Embrik Strand, 3, 1–50.
  17. Denis, J. (1939) Description d’un Zodarion nouveau du Portugal. Bulletin de la Société entomologique de France, 44, 89–92.
  18. https://doi.org/10.3406/bsef.1939.15363
  19. Denis, J. (1957) Zoologisch-systematische Ergebnisse der Studien-reise von H. Janetschek und W. Steiner in die Spanische Sierra Nevada. VII. Araneae. Sitzungsberichte der Österreichischen Akademie der Wissenschaften, 166, 265–302.
  20. Drensky, P. (1940) Die Spinnenfauna Bulgariens. IV. Unterordnung Arachnomorphae, II Gruppe Trionichia, Familien: Zodariidae, Dictynidae und Amaurobiidae. Mitteilungen aus den Königlichen Naturwissenschaftlichen Instituten in Sofia, 13, 169–194.
  21. Gaymard, M. & Lecigne, S. (2018) Contribution à la connaissance de l’aranéofaune (Araneae) du Gard et en particulier du massif des Gorges du Gardon (Occitanie, France). Bulletin de l’Association Française d’Arachnologie, 1, 1–39.
  22. Gaymard M., Mazzia, C., Danflous, S., Capowiez, Y., Cornic, J-F. & Lecigne, S. (2018) Découverte en France de Zodarion styliferum (Simon, 1870) (Araneae, Zodariidae). Revue Arachnologique, Série 2, 18–21.
  23. 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. Systematic biology, 59, 307–321. https://doi.org/10.1093/sysbio/syq010
  24. Hebert, P.D.N., Cywinska, A., Ball, S.L. & de Waard, J.R. (2003) Biological identifications through DNA barcodes. Proceedings of the Royal Society B: Biological Sciences, 270, 313–321.
  25. https://doi.org/10.1098/rspb.2002.2218
  26. Hedin, 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.
  27. https://doi.org/10.1006/mpev.2000.0883
  28. 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
  29. ICZN (1999) International code of zoological nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
  30. Jocqué, R. (1991) A generic revision of the spider family Zodariidae (Araneae). Bulletin of the American Museum of Natural History, 201, 1–160.
  31. 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
  32. Katoh, K. & Standley, D.M. (2013) MAFFT multiple sequence alignment software v.7: improvements in performance and usability. Molecular biology and evolution, 30, 772–780. https://doi.org/10.1093/molbev/mst010
  33. Kownacki, A., Szarek-Gwiazda, E. & Woźnicka, O. (2015) The importance of scanning electron microscopy (SEM) in taxonomy and morphology of Chironomidae (Diptera). European Journal of Environmental Sciences, 5, 41–44.
  34. https://doi.org/10.14712/23361964.2015.75
  35. Kulczyński, W. (1899) Arachnoidea opera Rev. E. Schmitz collecta in insulis Maderianis et in insulis Selvages dictis. Rozprawy i Sprawozdania z Posiedzen Wydzialu Matematyczno Przyrodniczego Akademji Umiejetnosci, Krakow, 36, 319–461.
  36. Kulczyński, W. (1905) Araneae nonnullae in insulus Maderianis collectae a Rev. E. Schmitz. Bulletin de l’Academie des Sciences de Cracovie, 440, 460.
  37. 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 (6), 1547–1549. https://doi.org/10.1093/molbev/msy096
  38. Michálek, O., Kuhn-Nentwig, L. & Pekár, S. (2019) High specific efficiency of venom of two prey-specialized spiders. Toxins, 11 (12), 687. https://doi.org/10.3390/toxins11120687
  39. Meier, R., Blaimer, B.B., Buenaventura, E., Hartop, E., von Rintelen, T., Srivathsan, A. & Yeo, D. (2022) A re‐analysis of the data in Sharkey et al.’s (2021) minimalist revision reveals that BINs do not deserve names, but BOLD Systems needs a stronger commitment to open science. Cladistics, 38, 264–275. https://doi.org/10.1111/cla.12489
  40. Minh, B.Q., Schmidt, H.A., Chernomor, O., Schrempf, D., Woodhams, M.D., von 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
  41. Ortiz, D. & Francke, O.F. (2015) Two new species of Bonnetina tarantulas (Theraphosidae: Theraphosinae) from Mexico: contributions to morphological nomenclature and molecular characterization of types. Journal of Natural History, 49, 685–707. https://doi.org/10.1080/00222933.2014.924770
  42. Ortiz, D., Pekár, S., Bilat, J. & Alvaraz, N. (2020) Poor performance of DNA barcoding and the impact of RAD loci filtering on the species delimitation of an Iberian ant-eating spider. Molecular Phylogenetics and Evolution, 154, 106997. https://doi.org/10.1016/j.ympev.2020.106997
  43. Ortiz, D., Pekár, S. & Dianat, M. (2022) Phylogenomics and loci dropout patterns of deeply diverged Zodarion ant‐eating spiders suggest a high potential of RAD‐seq for genus‐level spider phylogenetics. Cladistics, 38, 320–334. https://doi.org/10.1111/cla.12493
  44. Ortiz, D., Pekár, S. & Bryjová, A. (2023a) Gene flow assessment helps to distinguish strong genomic structure from speciation in an Iberian ant-eating spider. Molecular Phylogenetics and Evolution, 180, 107682. https://doi.org/10.1016/j.ympev.2022.107682
  45. Ortiz, D., Pekár, S., Bilat, J., Shafaie, S., Alvarez, N. & Gauthier, J. (2023b) Dynamic evolution of size and colour in the highly specialized Zodarion ant-eating spiders. Proceedings of the Royal Society B, 290, 20230797. https://doi.org/10.1098/rspb.2023.0797
  46. Pekár, S. & Král, J. (2002) Mimicry complex in two central European zodariid spiders (Araneae: Zodariidae): how Zodarion deceives ants. Biological Journal of the Linnean Society, 75, 517–532. https://doi.org/10.1046/j.1095-8312.2002.00043.x
  47. Pekár, S., Cardoso, P. & Meierrose, C. (2003) Additions to the knowledge of Portuguese zodariid spiders (Araneae: Zodariidae). Bulletin-British Arachnological Society, 12, 385–395.
  48. Pekár, S. (2004) Predatory behavior of two European ant-eating spiders (Araneae, Zodariidae). Journal of Arachnology, 32, 31–41. https://doi.org/10.1636/S02-15
  49. Pekár, S. & Cardoso, P. (2005) Ant-eating spiders (Araneae: Zodariidae) of Portugal: additions to the current knowledge. Zootaxa, 1009 (1), 51–60. https://doi.org/10.11646/zootaxa.1009.1.5
  50. Pekár, S., Král, J. & Lubin, Y. (2005a) Natural history and karyotype of some ant-eating zodariid spiders (Araneae, Zodariidae) from Israel. Journal of Arachnology, 33, 50–62. https://doi.org/10.1636/S03-2
  51. Pekár, S., Král, J., Malten, A. & Komposch, C. (2005b) Comparison of natural histories and karyotypes of two closely related ant-eating zodariid spiders, Zodarion hamatum and Z. italicum (Araneae, Zodariidae). Journal of Natural History, 39, 1583–1596.
  52. https://doi.org/10.1080/00222930400016762
  53. Pekár, S. & Toft, S. (2009) Can ant-eating Zodarion spiders (Araneae, Zodariidae) develop on a diet optimal for euryphagous arthropod predators? Physiological Entomology, 34, 195–201. https://doi.org/10.1111/j.1365-3032.2009.00672.x
  54. Pekár, S., Cardoso, P., Barriga, J.C. & Carvalho, J.C. (2011) Update to the zodariid spider fauna of the Iberian Peninsula and Madeira (Araneae: Zodariidae). Zootaxa, 2814 (1), 19–32. https://doi.org/10.11646/zootaxa.2814.1.2
  55. Pekár, S., Šmerda, J., Hrušková, M., Šedo, O., Muster, C., Cardoso, P., Zdráhal, Z., Korenko, S., Bureš, P., Líznarová, E. & Sentenská, L. (2012) Prey-race drives differentiation of biotypes in ant-eating spiders. Journal of Animal Ecology, 81 (4), 838–848.
  56. https://doi.org/10.1111/j.1365-2656.2012.01957.x
  57. Pekár, S., Šedo, O., Líznarová, E., Korenko, S. & Zdráhal, Z. (2014) David and Goliath: potent venom of an ant‐eating spider (Araneae) enables capture of a giant prey. Naturwissenschaften, 101 (7), 533–540. https://doi.org/10.1007/s00114-014-1189-8
  58. Pekár, S., Petráková, L., Šedo, O., Korenko, S. & Zdráhal, Z. (2018) Trophic niche, capture efficiency and venom profiles of six sympatric ant‐eating spider species (Araneae: Zodariidae). Molecular Ecology, 27, 1053–1064. https://doi.org/10.1111/mec.14485
  59. Pekár, S., Ortiz, D., Sentenská, L. & Šedo, O. (2022) Ecological specialization and reproductive isolation among closely related sympatric ant-eating spiders. Journal of Animal Ecology, 91 (9), 1855–1868. https://doi.org/10.1111/1365-2656.13767
  60. Shafaie, S., Mirshamsi, O., Aliabadian, M., Moradmand, M. & Marusik, Y. (2018) A new Pardosa species from northern Iran (Araneae, Lycosidae). Zootaxa, 4387 (2), 350–364. https://doi.org/10.11646/zootaxa.4387.2.6
  61. Shafaie, S. & Pekár, S. (2024) First record of Zodarion from the spinibarbe group in Spain (Araneae: Zodariidae). Serket, 20 (3), 275–280.
  62. Simon, E. (1870) Sur les aranéides de la famille des Enydes qui habitant l’Espagne et le Maroc. Revue et Magasin de Zoologie Pure Appliqée, 2 (22), 51–54 + 97–103 + 142–148.
  63. Simon, E. (1873) Aranéides nouveaux ou peu connus du midi de l’Europe. Extrait des Mémoires de la Société royale des Sciences de Liège, 2 (5), 187–351. [separately paginated version (pp. 1–174) seen and cited here.]
  64. https://doi.org/10.5962/bhl.title.124166
  65. Simon, E. (1874) Les Arachnides de France. Librarie Encyclopédique de Roter, tome premier. Roret, Paris. [unknown pagination]
  66. Simon, E. (1890) Etudes arachnologiques. 22e Mémoire. XXXIV. Etude sur les arachnides de l’Yemen. Annales de la Société Entomologique de France, 6 (10), 77–124.
  67. Simon, E. (1893) Histoire naturelle des araignées. Tome Premier. Deuxième Édition. Roret, Paris, 3 + 1084 pp. https://doi.org/10.5962/bhl.title.51973
  68. Simon, E. (1914) Les Arachnides de France. Synopsis générale et catalogue des espèces françaises de l’ordre des Araneae. Tome VI. Roret, Paris. [unknown pagination]
  69. Thorell, T. (1881) Studi sui Ragni Malesi e Papuani. III. Ragni dell’Austro Malesia e del Capo York, conservati nel Museo civico di storia naturale di Genova. Annali del Museo Civico di Storia Naturale di Genova, 17, 1–720.
  70. Walckenaer, C.A. (1826) Aranéides. In: Faune française ou histoire naturelle générale et particulière des animaux qui se trouvent en France, constamment ou passagèrement, à la surface du sol, dans les eaux qui le baignent et dans le littoral des mers qui le bornent par Viellot, Desmarrey, Ducrotoy, Audinet, Lepelletier et Walckenaer: Livraisou, Paris, pp. 11–12 + 1–96.
  71. World Spider Catalog (2025) World Spider Catalog. Version 25.5. Natural History Museum Bern, Bern. Available from http://wsc.nmbe.ch (accessed 15 January 2025) https://doi.org/10.24436/2
  72. Wunderlich, J. (2022) Some spiders (Araneae) of the Western Palearctic. Beiträge zur Araneologie, 15, 4–199.
  73. Wunderlich, J. (2023) Contribution to the spider (Araneida) fauna of the Algarve, Portugal, with notes on the phylogeny of the superfamily Araneoidea and the description of the new araneoid family Fonteferridae. Beiträge zur Araneologie, 16, 4–112.
  74. Zamani, A., Fric, Z.F., Gante, H.F., Hopkins, T., Orfinger, A.B., Scherz, M.D., Bartoňová, A.S. & Pos, D.D. (2022) DNA barcodes on their own are not enough to describe a species. Systematic Entomology, 47, 385–389. https://doi.org/10.1111/syen.12538

How to Cite

Shafaie, S., Pekár, S. & Ortiz, D. (2025) Integrative taxonomy of the Iberian Zodarion species of the rubidum and styliferum groups (Araneae: Zodariidae). Zootaxa, 5624 (1), 1–69. https://doi.org/10.11646/zootaxa.5624.1.1