Twigs that are not twigs: phylogenetic placement of crab spiders of the genus Tmarus of Sri Lanka with comments on the higher-level phylogeny of Thomisidae
Ilesha S. Ileperuma Arachchi A and Suresh P. Benjamin
A B C
+ Author Affiliations
- Author Affiliations
A National Institute of Fundamental Studies, Hantana Road, Kandy, Sri Lanka.
B Zoological Research Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany.
C Corresponding author. Email: suresh.benjamin@gmail.com
Invertebrate Systematics 33(3) 575-595 https://doi.org/10.1071/IS18074
Submitted: 12 September 2018 Accepted: 22 January 2019 Published: 4 June 2019
Abstract
Sri Lankan species of the genus Tmarus are cryptic spiders that live either on greenish moss-covered twigs or brownish dried-out twigs. They have remained taxonomically unrevised. The only known species was misplaced in Peritraeus Simon, 1895, which has never been subjected to phylogenetic evaluation. The present study is designed to investigate the phylogenetic placement of Sri Lankan Tmarus within Thomisidae as well as to assess the validity of Peritraeus and its relationship to Tmarus and Monaeses, if any. Using a multilocus molecular dataset (16S–ND1, 28S, CO1 and H3) we provide evidence that Tmarus and Monaeses are paraphyletic and that Peritraeus is a junior subjective synonym of Tmarus. Four species of Tmarus, including three new species, are now recorded from Sri Lanka. The legacy groups Alcimochthae and Tmarae are confirmed to be polyphyletic. Further, the following new species are described: Tmarus hiyarensis, sp. nov., T. viridomaculatus, sp. nov. and T. manojkaushalyai sp. nov. The following new combination is proposed: Tmarus hystrix (Simon, 1895), comb. nov. (this results in a homonym with Tmarus hystrix Caporiacco, 1954: we rename the species Tmarus caporioccoi Ileperuma Arachchi & Benjamin, replacement name).
Lsid: urn:lsid:zoobank.org:pub:05C2F204-3C31-43A6-8D7E-9AC56BFF1F6B
Additional keywords: biodiversity, cryptic behaviour, evolution, hotspot, laterigrade, mimicry, systematics
References
Álvarez-Padilla, F., and Hormiga, G. (2007). A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy. The Journal of Arachnology 35, 538–542.| A protocol for digesting internal soft tissues and mounting spiders for scanning electron microscopy.Crossref | GoogleScholarGoogle Scholar |
Astrin, J. J., Höfer, H., Spelda, J., Holstein, J., Bayer, S., Hendrich, L., Huber, B. A., Kielhorn, K-H., Krammer, H.J., Lemke, M., Monje, J. C., Moriniàre, J., Rulik, B., Petersen, M., Janssen, H., and Muster, C. (2016). Towards a DNA barcode reference database for spiders and harvestmen of Germany. PLoS One 11, e0162624.
| Towards a DNA barcode reference database for spiders and harvestmen of Germany.Crossref | GoogleScholarGoogle Scholar | 27681175PubMed |
Benjamin, S. P. (2000). Epidius parvati sp. n., a new species of the genus Epidius from Sri Lanka (Araneae: Thomisidae). Bulletin of the British Arachnological Society 11, 284–288.
Benjamin, S. P. (2001). The genus Oxytate L. Koch 1878 from Sri Lanka, with description of Oxytate taprobane sp. n. (Araneae: Thomisidae). Journal of South Asian Natural History 5, 153–158.
Benjamin, S. P. (2004). Taxonomic revision and phylogenetic hypothesis for the jumping spider subfamily Ballinae (Araneae, Salticidae). Zoological Journal of the Linnean Society 142, 1–82.
| Taxonomic revision and phylogenetic hypothesis for the jumping spider subfamily Ballinae (Araneae, Salticidae).Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P. (2008). Family ties: molecular phylogeny of crab spiders. Cladistics 24, 708–722.
| Family ties: molecular phylogeny of crab spiders.Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P. (2011). Phylogenetics and comparative morphology of crab spiders (Araneae: Dionycha, Thomisidae). Zootaxa 3080, 1–108.
Benjamin, S. P. (2016). Revision of Cebrenninus Simon, 1887 with description of one new genus and six new species (Araneae: Thomisidae). Revue Suisse de Zoologie 123, 179–200.
| Revision of Cebrenninus Simon, 1887 with description of one new genus and six new species (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P., and Jalee, Z. (2007). Redescription of Tagulis mystacinus (Araneae: Thomisidae) from Sri Lanka. Zootaxa 1560, 63–68.
| Redescription of Tagulis mystacinus (Araneae: Thomisidae) from Sri Lanka.Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P., and Jaleel, Z. (2010). The genera Haplotmarus Simon, 1909 and Indoxysticus gen. nov.: two enigmatic genera of crab spiders from the Oriental region (Araneae: Thomisidae). Revue Suisse de Zoologie 117, 159–167.
| The genera Haplotmarus Simon, 1909 and Indoxysticus gen. nov.: two enigmatic genera of crab spiders from the Oriental region (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P., Dimitrov, D., Gillespie, R. G., and Hormiga, G. (2008). Family ties: molecular phylogeny of crab spiders (Araneae: Thomisidae). Cladistics 24, 708–722.
| Family ties: molecular phylogeny of crab spiders (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Benjamin, S. P., Ileperuma Arachchi, I. S., and Herath, B. P. (2016). First record of Platythomisus sudeepi from Sri Lanka (Araneae: Thomisidae). Arachnology 17, 165–168.
| First record of Platythomisus sudeepi from Sri Lanka (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Boc, A., Diallo, A. B., and Makarenkov, V. (2012). T-REX: a web server for inferring, validating and visualizing phylogenetic trees and networks. Nucleic Acids Research 40, W573–W579.
| T-REX: a web server for inferring, validating and visualizing phylogenetic trees and networks.Crossref | GoogleScholarGoogle Scholar | 22675075PubMed |
Bremer, K. (1994). Branch support and tree stability. Cladistics 10, 295–304.
| Branch support and tree stability.Crossref | GoogleScholarGoogle Scholar |
Caporiacco, L. (1954). Araignées de la Guyane Française du Muséum d’Histoire Naturelle de Paris. Commentationes Pontificia Academia Scientiarum 16, 45–193.
Castresana, J. (2000). Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17, 540–552.
| Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis.Crossref | GoogleScholarGoogle Scholar | 10742046PubMed |
Chickering, A. M. (1965a). Panamanian spiders of the genus Tmarus (Araneae, Thomisidae). Bulletin of the Museum of Comparative Zoology 133, 337–368.
Chickering, A. M. (1965b). Five new species of the genus Tmarus (Araneae, Thomisidae) from the West Indies. Psyche 72, 229–240.
| Five new species of the genus Tmarus (Araneae, Thomisidae) from the West Indies.Crossref | GoogleScholarGoogle Scholar |
Dippenaar-Schoeman, A. (1984). The crab-spiders of southern Africa (Araneae: Thomisidae). 4. The genus Monaeses Thorell 1869. Phytophylactica 16, 101–116.
Dippenaar-Schoeman, A. S. (1985). The crab-spiders of southern Africa (Araneae: Thomisidae). 5. The genus Tmarus Simon, 1875. Phytophylactica 17, 115–128.
Edgecombe, G. D., and Giribet, G. (2006). A century later – a total evidence re-evaluation of the phylogeny of scutigeromorph centipedes (Myriapoda: Chilopoda). Invertebrate Systematics 20, 503–525.
| A century later – a total evidence re-evaluation of the phylogeny of scutigeromorph centipedes (Myriapoda: Chilopoda).Crossref | GoogleScholarGoogle Scholar |
Garb, J. E., and Gillespie, R. G. (2009). Diversity despite dispersal: colonization history and phylogeography of Hawaiian crab spiders inferred from multilocus genetic data. Molecular Ecology 18, 1746–1764.
| Diversity despite dispersal: colonization history and phylogeography of Hawaiian crab spiders inferred from multilocus genetic data.Crossref | GoogleScholarGoogle Scholar | 19302468PubMed |
Gawryszewski, F. M., Calero-Torralbo, M. A., Gillespie, R. G., Rodríguez-Gironés, M. A., and Herberstein, M. E. (2017). Correlated evolution between coloration and ambush site in predators with visual prey lures. Evolution 71, 2010–2021.
| Correlated evolution between coloration and ambush site in predators with visual prey lures.Crossref | GoogleScholarGoogle Scholar | 28543028PubMed |
Goloboff, P. A., Farris, J. S., Kallersjo, M., Oxelman, B., Ramırez, M. J., and Szumik, C. A. (2003). Improvements to resampling measures of group support. Cladistics 19, 324–332.
| Improvements to resampling measures of group support.Crossref | GoogleScholarGoogle Scholar |
Goloboff, P. A., Farris, J. S., and Nixon, K. C. (2008). TNT, a free program for phylogenetic analysis. Cladistics 24, 774–786.
| TNT, a free program for phylogenetic analysis.Crossref | GoogleScholarGoogle Scholar |
Kim, S. T., and Lee, S. Y. (2012). Arthropoda: Arachnida: Araneae: Thomisidae. Thomisid spiders. Invertebrate Fauna of Korea 21, 1–88.
Machado, M., Teixeira, R. A., and Lise, A. A. (2017). Cladistic analysis supports the monophyly of the Neotropical crab spider genus Epicadus and its senior synonymy over Tobias (Araneae: Thomisidae). Invertebrate Systematics 31, 442–455.
| Cladistic analysis supports the monophyly of the Neotropical crab spider genus Epicadus and its senior synonymy over Tobias (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Maddison, W. P., and Maddison, D. R. (2009). Mesquite: a modular system for evolutionary analysis. Version 2.72. Available at: http://mesquiteproject.org [accessed 21 February 2019].
Metcalfe, N. B., Valdimarsson, S. K., and Fraser, N. H. C. (1997). Habitat profitability and choice in a sit-and-wait predator: juvenile salmon prefer slower currents on darker nights. Journal of Animal Ecology 66, 866–875.
| Habitat profitability and choice in a sit-and-wait predator: juvenile salmon prefer slower currents on darker nights.Crossref | GoogleScholarGoogle Scholar |
Moradmand, M., Schönhofer, A. L., and 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.
| Molecular phylogeny of the spider family Sparassidae with focus on the genus Eusparassus and notes on the RTA-clade and ‘Laterigradae’.Crossref | GoogleScholarGoogle Scholar | 24508702PubMed |
Ono, H. (1977). Thomisidae aus Japan I. Das Genus Tmarus Simon (Arachnida: Araneae). Acta Arachnologica 27, 61–84.
Ono, H. (1988). A revisional study of the spider family Thomisidae (Arachnida, Araneae) of Japan. National Science Museum,Tokyo ii, 1–252.
Polotow, D., Carmichael, A., and Griswold, C. E. (2015). Total evidence analysis of the phylogenetic relationships of Lycosoidea spiders (Araneae, Entelegynae). Invertebrate Systematics 29, 124–163.
| Total evidence analysis of the phylogenetic relationships of Lycosoidea spiders (Araneae, Entelegynae).Crossref | GoogleScholarGoogle Scholar |
Ramírez, M. J. (2014). The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History 390, 1–374.
| The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae).Crossref | GoogleScholarGoogle Scholar |
Sanger, F., Nicklen, S., and Coulson, A. R. (1977). DNA sequencing with chain-terminating inhibitors. Proceedings of the National Academy of Sciences of the United States of America 74, 5463–5467.
| DNA sequencing with chain-terminating inhibitors.Crossref | GoogleScholarGoogle Scholar | 271968PubMed |
Sen, S., Dhali, D. C., Saha, S., and Raychaudhuri, D. (2015). Spiders (Araneae: Arachnida) of Reserve Forests of Dooars: Gorumara National Park, Chapramari Wildlife Sanctuary and Mahananda Wildlife Sanctuary. World Scientific News 20, 1–339.
Silva-Moreira, T., and Machado, M. (2016). Taxonomic revision of the crab spider genus Epicadus Simon, 1895 (Arachnida: Araneae: Thomisidae) with notes on related genera of Stephanopinae Simon, 1895. Zootaxa 4147, 281–310.
| Taxonomic revision of the crab spider genus Epicadus Simon, 1895 (Arachnida: Araneae: Thomisidae) with notes on related genera of Stephanopinae Simon, 1895.Crossref | GoogleScholarGoogle Scholar | 27515619PubMed |
Simon, E. (1875). ‘Les Arachnides de France.’ Paris 2, 1–350.
Simon, E. (1895). ‘Histoire Naturelle des Araignées.’ Paris 1, 761–1084.
Sirvid, P. J., Moore, N. E., Chambers, G. K., and Prendergast, K. (2013). A preliminary molecular analysis of phylogenetic and biogeographic relationships of New Zealand Thomisidae (Araneae) using a multi-locus approach. Invertebrate Systematics 27, 655–672.
| A preliminary molecular analysis of phylogenetic and biogeographic relationships of New Zealand Thomisidae (Araneae) using a multi-locus approach.Crossref | GoogleScholarGoogle Scholar |
Stamatakis, A. (2006). RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics 22, 2688–2690.
| RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models.Crossref | GoogleScholarGoogle Scholar | 16928733PubMed |
Talavera, G., and Castresana, J. (2007). Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments. Systematic Biology 56, 564–577.
| Improvement of phylogenies after removing divergent and ambiguously aligned blocks from protein sequence alignments.Crossref | GoogleScholarGoogle Scholar | 17654362PubMed |
Tang, G., and Li, S. Q. (2009). The crab spiders of th genus Tmarus from Xishuangbanna, Yunnan, China (Araneae: Thomisidae). Zootaxa 2223, 48–68.
| The crab spiders of th genus Tmarus from Xishuangbanna, Yunnan, China (Araneae: Thomisidae).Crossref | GoogleScholarGoogle Scholar |
Thorell, T. (1869). On European spiders. Part I. Review of the European genera of spiders, preceded by some observations on zoological nomenclature. Nova Acta Regiae Societatis Scientiarum Upsaliensis 3, 1–108.
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, L. R., Benjamin, S. P., Bond, J. E., Grismado, C. J., Hasanf, 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., Szúts, T., Ubick, D., Vink, C. J., Wood, H. M., and 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.
| The spider tree of life: phylogeny of Araneae based on target-gene analyses from an extensive taxon sampling.Crossref | GoogleScholarGoogle Scholar |
World Spider Catalog (2018). World Spider Catalog. Natural History Museum Bern. Version 19.0. Available at: http://wsc.nmbe.ch [accessed January 2018].
Yang, Z. Z., Zhu, M.-S., and Song, D. X. (2005). Two new species of the spider genus Tmarus Simon 1875 (Araneae: Thomisidae) from China. Acta Arachnologica 54, 95–98.
| Two new species of the spider genus Tmarus Simon 1875 (Araneae: Thomisidae) from China.Crossref | GoogleScholarGoogle Scholar |
Zamani, A. (2015). The spider collection (Arachnida: Araneae) of the Zoological Museum of the Iranian Research Institute of Plant Protection, with new species records for Iran. Arachnologische Mitteilungen 50, 11–18.
| The spider collection (Arachnida: Araneae) of the Zoological Museum of the Iranian Research Institute of Plant Protection, with new species records for Iran.Crossref | GoogleScholarGoogle Scholar |
