Taxonomic revision and cladistic analysis of ghost spiders of the genus Tafana Simon, 1903 (Araneae: Dionycha, Anyphaenidae), with the descriptions of twelve new species
Abstract
The diversity of the genus Tafana Simon, 1903 is poorly known in the Neotropical regions. In this work we provide a taxonomic review of the genus as well as a phylogenetic analysis. The ingroup of the analysis is composed of sixteen species of Tafana and the outgroup is composed of five representatives of Anyphaenidae. The sister-group recovered for Tafana is the clade Aysha + Xiruana, being supported by the embolic process on the male bulb. Two species groups within Tafana are herein proposed, the silhavyi group and the riveti group, based on two exclusive synapomorphies in the male bulb. We redescribe Tafana quelchi and present a description of the previously unknown female of Tafana silhavyi, both from Venezuela. In addition, we describe the first adult specimens of Tafana straminea. Twelve new species, along with several previously described species, are described, illustrated and mapped: T. riveti, T. straminea, T. quelchi, T. kunturmarqa sp. nov., T. humahuaca sp. nov., T. pastaza sp. nov., T. nevada sp. nov., T. huatanay sp. nov. and T. ruizi sp. nov. from the riveti species group; T. maracay sp. nov., T. arawak sp. nov., T. chimire sp. nov. and T. pitieri sp. nov. from the silhavyi species group; T. oliviae sp. nov. from Argentina and T. orinoco sp. nov. from Venezuela, neither of which belongs to any species group. We also discuss the genital morphology of the species groups based on the results of the phylogenetic analysis. Furthermore, distribution maps for all species, including new records for T. riveti, T. straminea and T. quelchi, are presented.
References
Azevedo G.H.F., Griswold C.E. & Santos A.J. 2018. Systematics and evolution of ground spiders revisited (Araneae, Dionycha, Gnaphosidae). Cladistics 34: 579–626. https://doi.org/10.1111/cla.12226
Berland L. 1913. Araignées. In: Service géographique de l’Armée. Mission du Service géographique de l’Armée pour la mesure d’un arc du méridien équatorial en Amérique du Sud (1899–1906) 10: 78–119. Gauthier-Villars, Paris. https://doi.org/10.5962/bhl.title.980
Bonaldo A.B. 2000. Taxonomia da subfamília Corinninae (Araneae, Corinnidae) nas regiões Neotropical e Neártica. Iheringia 89: 3–148.
Bonnet P. 1955. Bibliographia araneorum. Analyse méthodique de toute la littérature aranéologique jusqu’en 1939. Tome II. Systématique des araignées (Étude par ordre alphabétique) [1e partie: A-B]: 1–918. Douladoure, Toulouse.
Bonnet P. 1959. Bibliographia araneorum. Analyse méthodique de toute la littérature aranéologique jusqu’en 1939. Tome II. Systématique des araignées (Étude par ordre alphabétique) (5e partie: T-Z): 4231–5058. Douladoure, Toulouse.
Brescovit A.D. 1991a. Revalidação do gênero Isigonia Simon, com descrição de uma espécie nova (Araneae, Anyphaenidae). Revista Brasileira de Entomologia 35 (4): 721–727.
Brescovit A.D. 1991b. Hibana, novo gênero de aranhas da família Anyphaenidae (Arachnida, Araneae). Revista Brasileira de Entomologia 35 (4): 729–744.
Brescovit A.D. 1992a. Revisão das aranhas do gênero Macrophyes O. Pickard-Cambridge, da região Neotropical (Araneae, Anyphaenidae). Revista Brasileira de Entomologia 36 101–106.
Brescovit A.D. 1992b. Revisão do grupo prospera do gênero Aysha Keyserling, 1891 na região Neotropical (Araneae, Anyphaenidae). Iheringia Série Zoológica: 72: 23–104.
Brescovit A.D. 1993a. Novas espécies do gênero Macrophyes O.P.-Cambridge da região Neotropical (Araneae, Anyphaenidae). Iheringia 75: 113–116.
Brescovit A.D. 1993b. Thaloe e Bromelina, novos gêneros de aranhas neotropicais da família Anyphaenidae (Arachnida, Araneae). Revista Brasileira de Entomologia 37: 693–703.
Brescovit A.D. 1997. Revisão de Anyphaeninae Bertkau a nível de gêneros na região Neotropical (Araneae, Anyphaenidae). Revista Brasileira de Zoologia 13: 1–187. https://doi.org/10.1590/S0101-81751996000500001
Brescovit A.D. 1999. Revisão das aranhas do gênero Lupettiana Brescovit (Araneae, Anyphaenidae, Anyphaeninae). Revista Brasileira de Zoologia 16: 63–76. https://doi.org/10.1590/S0101-81751999000600008
Brescovit A.D. & Oliveira L.F.M. 2019. Three new species of Thaloe Brescovit (Araneae: Anyphaenidae, Anyphaeninae), with descriptions of the first known females of the genus. Zootaxa 4624 (2): 256–266. https://doi.org/10.11646/zootaxa.4624.2.8
Bremer K. 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution 42: 795–803.
Bremer K. 1994. Branch support and tree stability. Cladistics 10: 295–304.
Cabra-García J. & Brescovit A.D. 2016. Revision and phylogenetic analysis of the orb-weaving spider genus Glenognatha Simon, 1887 (Araneae, Tetragnathidae). Zootaxa 4069 (1): 1–183. https://doi.org/10.11646/zootaxa.4069.1.1
Caporiacco L. Di. 1955. Estudios sobre los aracnidos de Venezuela 2a parte: Araneae. Acta biologica Venezuelica 1 (16): 245–448.
Ceccarelli F.S., Opell B.D., Haddad C.R., Raven R.J., Soto E.M. & Ramírez M.J. 2016. Around the world in eight million years: historical biogeography and evolution of the spray zone spider Amaurobioides (Araneae: Anyphaenidae). PLoS One 11 (10): e0163740. https://doi.org/10.1371/journal.pone.0163740
Ceccarelli F.S., Koch M.N., Soto E.M., Barone M.L., Arnedo M.A. & Ramírez M.J. 2019. The grass was greener: repeated evolution of specialized morphologies and habitat shifts in ghost spiders following grassland expansion in South America. Systematics Biology 68 (1): 63–77. https://doi.org/10.1093/sysbio/syy028
Coddington J.A. 1990. Ontogeny and homology in the male palpus of orb-weaving spiders and their relatives, with comments on phylogeny (Araneoclada: Araneoidea, Deinopoidea). Smithsonian Contributions to Zoology 496: 1–52.
Dupérré N. & Tapia E. 2016. Overview of the Anyphaenids (Araneae, Anyphaeninae, Anyphaenidae) spider fauna from the Chocó forest of Ecuador, with the description of thirteen new species. European Journal of Taxonomy 255: 1–50. https://doi.org/10.5852/ejt.2016.255
Fitch W.M. 1971. Toward defining the course of evolution: minimum change for a specific tree topology. Systematic Zoology 20 (4): 404–416.
Goodman M., Olson C.B., Beeber J.E. & Czelusniak J. 1982. New perspectives in the molecular biological analysis of mammalian phylogeny. Acta Zoologica Fennica 169: 19–35.
Goloboff P.A. 2008. Calculating SPR distances between trees. Cladistics 24: 591–597.
Goloboff P.A. & Catalano S.A. 2016. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics 32: 1–18.
Goloboff P. & Farris J. 2001. Methods for quick consensus estimation. Cladistics 17: S26–S34. https://doi.org/10.1006/clad.2000.0156
Goloboff P.A., Farris J.S., Källersjö M., Oxelman B., Ramírez M.J. & Szumik C.A. 2003. Improvements to resampling measures of group support. Cladistics 19: 324–332.
Goloboff P.A., Farris S. & Nixon K. 2008. TNT, a free program for phylogenetic analysis. Cladistics 24: 774–786.
Gonzalez Marquez M.E. & Ramírez M.J. 2012. A revision and phylogenetic analysis of the spider genus Aysenia Tullgren (Araneae: Anyphaenidae, Amaurobioidinae). Zootaxa 3201 (1): 1–26.
https://doi.org/10.11646/zootaxa.3201.1.1
Griswold C.E., Coddington J.A., Hormiga G. & Scharff N. 1998. Phylogeny of the orb-web building spiders (Araneae, Orbiculariae: Deinopoidea, Araneoidea). Zoological Journal of the Linnean Society 123 (1): 1–99. https://doi.org/10.1111/j.1096-3642.1998.tb01290.x
Izquierdo M.A. & Ramırez M.J. 2008. Two new spider species of the genera Aysenia and Aysenoides from southern Chile and Argentina: description and phylogenetic relationships (Araneae: Anyphaenidae, Amaurobioidinae). Zootaxa 1861 (1): 29–43. https://doi.org/10.11646/zootaxa.1861.1.3
Järvi T.H. 1912. Das Vaginalsystem der Sparassiden. Annales Academiae Scientiarum Fennicae (A) 4: 1–131.
Koch L. 1866. Die Arachniden – Familie der Drassiden. Parts 1–6: 1–304.
Labarque F.M., Soto E.M., Ramírez M.J. & Arnedo M.A. 2015. Chasing ghosts: the phylogeny of Amaurobioidinae ghost spiders (Araneae, Anyphaenidae). Zoologica Scripta 44 (5): 550–561.
Laborda A., Ramırez M.J. & Pizarro-Araya J. 2013. New species of the spider genera Aysenia and Aysenoides from Chile and Argentina: description and phylogenetic relationships (Araneae: Anyphaenidae, Amaurobioidinae). Zootaxa 3731: 133–152. https://doi.org/10.11646/zootaxa.3731.1.6
Nixon K.C. 2002. Winclada, ver. 1.00.08. Published by the author, Ithaca, NY.
Oliveira L.F.M. & Brescovit A.D. 2015a. A taxonomic revision of the Neotropical spider genus Xiruana Brescovit 1997 (Araneae: Anyphaenidae, Anyphaeninae). Zootaxa 3980 (2): 201–229. https://doi.org/10.11646/zootaxa.3980.2.3
Oliveira L.F.M. & Brescovit A.D. 2015b. Two new species of the Neotropical spider genus Umuara Brescovit (Araneae, Anyphaenidae, Anyphaeninae) from Brazil. Zootaxa 3980 (3): 435–441. https://doi.org/10.11646/zootaxa.3980.3.7
Oliveira L.F.M. & Brescovit A.D. 2016. The first male of the spider genus Temnida Simon, 1896 (Araneae: Anyphaenidae: Anyphaeninae). Arachnology 17 (1): 47–51. https://doi.org/10.13156/arac.2006.17.1.47
Petrunkevitch A. 1911. A synonymic index-catalogue of spiders of North, Central and South America with all adjacent islands, Greenland, Bermuda, West Indies, Terra del Fuego, Galapagos, etc. Bulletin of the American Museum of Natural History 29: 1–791.
Petrunkevitch A. 1928. Systema Aranearum. Transactions of the Connecticut Academy of Arts and Sciences 29: 1–270.
Platnick N.I. & Gertsch W.J. 1976. The suborders of spiders: a cladistic analysis (Arachnida, Araneae). The American Museum of Natural History 2607: 1–15.
Pocock R.I. 1895. Description of two new species obtained by Messrs J.J. Quelch and F. Mc Connell on the summit of Mount Roraima, in Demerara, with a note upon the systematic position of the genus Desis. Annals and Magazine of Natural History 16 (6): 139–143. https://doi.org/10.1080/00222939508680241
Pocock R.I. 1900. Myriapoda and Arachnida. In: Report on a Collection made by M. M. F.V. McConnell and J.J. Quelch at Mount Roraima in British Guiana. Transactions of the Linnean Society of London 2 (8): 64–71. https://doi.org/10.1111/j.1096-3642.1900.tb00308.x
Ramírez M.J. 1995a. A phylogenetic analysis of the subfamilies of Anyphaenidae (Arachnida, Araneae). Entomologica Scandinavica 26: 361–384.
Ramírez M.J. 1995b. Revisión y filogenía del género Monapia, con notas sobre otras Amaurobioidinae (Araneae, Anyphaenidae). Boletin de la Sociedad de Biologia de Concepcion 66: 71–102.
Ramírez M.J. 1999. New species and cladistic reanalysis of the spider genus Monapia (Araneae: Anyphaenidae, Amaurobioidinae). Journal of Arachnology 27: 415–431.
Ramírez M.J. 2003. The spider subfamily Amaurobioidinae (Araneae: Anyphaenidae): A phylogenetic revision at generic level. Bulletin of the Museum of Natural History 277: 1–262. https://doi.org/fv3bxr
Ramírez M.J. 2007. Homology as a parsimony problem: a dynamic homology approach for morpho-logical data. Cladistics 23: 588–612. https://doi.org/10.1111/j.1096-0031.2007.00162.x
Ramírez M.J. 2014. The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390: 1–374. https://doi.org/10.1206/821.1
Ramírez M.J., Ansaldi M.J. & Puglisi A.F. 2004. Description of the females of Oxysoma itambezinho Ramírez and Monapia tandil Ramírez, and their effects on the generic relationships of Gayennini (Araneae, Anyphaenidae, Amaurobioidinae). Zootaxa 668 (1): 1–8. https://doi.org/10.11646/zootaxa.668.1.1
Roewer C.F. 1954. Katalog der Araneae von 1758 bis 1940. Vol. 2: 1–923. Royal Belgian Institute of Natural Sciences, Brussels.
Rubio G.D. & Ramírez M.J. 2015. Taxonomic revision of the American spider genus Arachosia (Araneae: Anyphaenidae). Zootaxa 3932 (1): 1–105. https://doi.org/10.11646/zootaxa.3932.1.1
Sereno P.C. 2007. Logical basis for morphological characters in phylogenetics. Cladistics 23: 565–587.
Sierwald P. 1989. Morphology and ontogeny of female copulatory organs in American Pisauridae, with special reference to homologous features (Arachnida: Araneae). Smithsonian Contributions to Zoology 484: 1–24.
Simon E. 1903. Descriptions d’Arachnides nouveaux. Annales de la Société entomologique de Belgique 47: 21–39.
Soto M.E. & Ramírez M.J. 2012. Revision and phylogenetic analysis of the spider genus Philisca Simon (Araneae: Anyphaenidae, Amaurobioidinae). Zootaxa 3443 (1): 1–65. https://doi.org/10.11646/zootaxa.3443.1.1
Soto M.E., Labarque M.F., Ceccarelli S.F., Arnedo A.M., Pizarro-Araya J. & Ramírez M. 2017. The life and adventures of an eight-legged castaway: colonization and diversification of Philisca ghost spiders on Robinson Crusoe Island (Araneae, Anyphaenidae). Molecular Phylogenetics and Evolution 107: 132–141. https://doi.org/10.1016/j.ympev.2016.10.017
Werenkraut V. & Ramírez M.J. 2009. A revision and phylogenetic analysis of the spider genus Coptoprepes Simon (Araneae: Anyphaenidae, Amaurobioidinae). Zootaxa 2212 (1): 1–40. https://doi.org/10.11646/zootaxa.2212.1.1
World Spider Catalog. 2020. World Spider Catalog. Version 20.0. Natural History Museum Bern. Available from http://wsc.nmbe.ch [accessed 26 Jun. 2020].
Copyright (c) 2021 Luiz Fernando M. De Oliveira, Antonio Domingos Brescovit
This work is licensed under a Creative Commons Attribution 4.0 International License.
Creative Commons Copyright Notices
Authors who publish with this journal agree to the following terms:
- Authors retain copyright and grant the journal right of first publication with the work simultaneously licensed under a Creative Commons Attribution License (CC BY 4.0) that allows others to share the work with an acknowledgement of the work's authorship and initial publication in this journal.
- Authors are able to enter into separate, additional contractual arrangements for the non-exclusive distribution of the journal's published version of the work (e.g., post it to an institutional repository or publish it in a book), with an acknowledgement of its initial publication in this journal.
- Authors are NOT ALLOWED TO post their work online (e.g., in institutional repositories or on their website) prior to and during the submission process, as it can lead to taxonomic issues.
