CSIRO Publishing blank image blank image blank image blank imageBooksblank image blank image blank image blank imageJournalsblank image blank image blank image blank imageAbout Usblank image blank image blank image blank imageShopping Cartblank image blank image blank image You are here: Journals > Invertebrate Systematics   
Invertebrate Systematics
Journal Banner
  Systematics, Phylogeny and Biogeography
blank image Search
blank image blank image
blank image
  Advanced Search

Journal Home
About the Journal
Editorial Structure
Current Issue
Just Accepted
All Issues
Special Issues
Virtual Issues
Sample Issue
For Authors
General Information
Submit Article
Author Instructions
Open Access
For Referees
Referee Guidelines
Review an Article
Annual Referee Index
For Subscribers
Subscription Prices
Customer Service
Print Publication Dates

blue arrow e-Alerts
blank image
Subscribe to our Email Alert or RSS feeds for the latest journal papers.

red arrow Connect with us
blank image
facebook twitter logo LinkedIn


Article << Previous     |     Next >>   Contents Vol 25(2)

The evolutionary and biogeographic history of the armoured harvestmen – Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida)

Prashant P. Sharma A B and Gonzalo Giribet A

A Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.
B Corresponding author. Email: psharma@fas.harvard.edu

Invertebrate Systematics 25(2) 106-142 http://dx.doi.org/10.1071/IS11002
Submitted: 11 January 2011  Accepted: 20 June 2011   Published: 30 September 2011

 Full Text
 PDF (5.7 MB)
 Supplementary Material
 Export Citation

We investigated the internal phylogeny of Laniatores, the most diverse suborder of Opiliones, using sequence data from 10 molecular loci: 12S rRNA, 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase subunit I (COI), cytochrome b, elongation factor-1α, histones H3 and H4, and U2 snRNA. Exemplars of all previously described families of Laniatores were included, in addition to two families – Petrobunidae, fam. nov. and Tithaeidae, fam. nov. – that we erect herein. Data analyses were based on maximum likelihood and Bayesian approaches on static alignments, and included phylogenetic tree estimation, molecular dating, and biogeographic analysis of ancestral area reconstruction. The results obtained include the monophyly of Laniatores and the infraorder Grassatores – the focus of this study – as well as support for numerous interfamilial relationships. The two new families described cluster with other South-east Asian families (Podoctidae and Epedanidae). Diversification of Laniatores is estimated at ~348 Mya, and origin of most Grassatores superfamilies occurs in a ~25 million year span of time immediately after the end-Permian mass extinction (254 Mya). Ancestral range reconstruction of the clade (Samooidea + Zalmoxoidea) suggests that the ancestral range of Samooidea comprises West Tropical Gondwana (West Africa + Neotropics), whereas that of Zalmoxoidea is exclusively Neotropical. The following additional taxonomic changes are proposed: (1) Remyus is transferred to Phalangodidae, and (2) Escadabiidae and Kimulidae are transferred to Zalmoxoidea.

Additional keywords: biogeography, Briggsidae, Grassatores, Guasiniidae, Insidiatores, molecular dating, Petrobunidae, Tithaeidae.


Castresana, J. (2000). Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17, 540–552.

Colgan, D. J., McLauchlan, A., Wislon, G. D. F., Livingston, S., Macaranas, J., Edgecombe, G. D., Cassis, G., and Gray, M. R. (1998). Molecular phylogenetics of the Arthropoda: relationships based on histone H3 and U2 snRNA DNA sequences. Australian Journal of Zoology 46, 419–437.
CrossRef |

DaSilva, M. B., and Gnaspini, P. (2009). A systematic revision of Goniosomatinae (Arachnida: Opiliones: Gonyleptidae), with a cladistic analysis and biogeographical notes. Invertebrate Systematics 23, 530–624.
CrossRef |

DaSilva, M. B., and Pinto-da-Rocha, R. (2010). Systematic review and cladistic analysis of the Hernandariinae (Opiliones: Gonyleptidae). Zoologia 27, 577–642.

Derkarabetian, S., Steinmann, D. B., and Hedin, M. (2010). Repeated and time-correlated morphological convergence in cave-dwelling harvestmen (Opiliones, Laniatores) from montane western North America. PLoS ONE 5, e10388.
CrossRef |

Drummond, A. J., and Rambaut, A. (2007). BEAST: Bayesian evolutionary analysis by sampling trees. BMC Evolutionary Biology 7, 214.
CrossRef |

Drummond, A. J., Ho, S. Y. W., Phillips, M. J., and Rambaut, A. (2006). Relaxed phylogenetics and dating with confidence. PLoS Biology 4, e88.
CrossRef |

Dunlop, J. A. (2007). Paleontology. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 247–265. (Harvard University Press: Cambridge, MA, USA.)

Dunlop, J. A., and Anderson, L. I. (2005). A fossil harvestman (Arachnida, Opiliones) from the Mississippian of East Kirkton, Scotland. The Journal of Arachnology 33, 482–489.
CrossRef |

Dunlop, J. A., Anderson, L. I., Kerp, H., and Hass, H. (2003). Preserved organs of Devonian harvestmen. Nature 425, 916.
CrossRef |

Dunlop, J. A., Anderson, L. I., Kerp, H., and Hass, H. (2004). A harvestman (Arachnida: Opiliones) from the Early Devonian Rhynie cherts, Aberdeenshire, Scotland. Transactions of the Royal Society of Edinburgh. Earth Sciences 94, 341–354.

Edgar, R. C. (2004). MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32, 1792–1797.
CrossRef |

Edgecombe, G. D., and Giribet, G. (2006). A century later– a total evidence re-evaluation of scutigeromorph centipedes (Myriapoda: Chilopoda). Invertebrate Systematics 20, 503–525.
CrossRef |

Edgecombe, G. D., Giribet, G., and Wheeler, W. C. (2002). Phylogeny of Henicopidae (Chilopoda: Lithobiomorpha): a combined analysis of morphology and five molecular loci. Systematic Entomology 27, 31–64.
CrossRef |

Folmer, O., Black, M., Hoeh, W., Lutz, R., and Vrijenhoek, R. C. (1994). DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metazoan invertebrates. Molecular Marine Biology and Biotechnology 3, 294–299.

Forster, R. R. (1954). The New Zealand harvestmen (sub-order Laniatores). Canterbury Museum Bulletin 2, 1–329.

Giribet, G., and Kury, A. B. (2007). Phylogeny and biogeography. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 62–87. (Harvard University Press: Cambridge, MA, USA.)

Giribet, G., Carranza, S., Baguñà, J., Riutort, M., and Ribera, C. (1996). First molecular evidence for the existence of a Tardigrada + Arthropoda clade. Molecular Biology and Evolution 13, 76–84.

Giribet, G., Rambla, M., Carranza, S., Baguñà, J., Riutort, M., and Ribera, C. (1999). Phylogeny of the arachnid order Opiliones (Arthropoda) inferred from a combined approach of complete 18S and partial 28S ribosomal DNA sequences and morphology. Molecular Phylogenetics and Evolution 11, 296–307.
CrossRef |

Giribet, G., Edgecombe, G. D., Wheeler, W. C., and Babbitt, C. (2002). Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data. Cladistics 18, 5–70.

Giribet, G., Vogt, L., Pérez González, A., Sharma, P., and Kury, A. B. (2010). A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores. Cladistics 26, 408–437.

Hara, M. R., Gnaspini, P., and Machado, G. (2003). Male egg guarding behavior in the Neotropical harvestman Ampheres leucopheus (Mello-Leitão 1922) (Opiliones, Gonyleptidae). The Journal of Arachnology 31, 441–444.
CrossRef |

Hedin, M., Derkarabetian, S., McCormack, M., and Richart, C. (2010). The phylogenetic utility of the nuclear protein-encoding gene EF-1α for resolving recent divergences in Opiliones, emphasizing intron evolution. The Journal of Arachnology 38, 9–20.
CrossRef |

Huelsenbeck, J. P. and Ronquist, F. (2005). Bayesian analyses of molecular evolution using MrBayes. In ‘Statistical Methods in Molecular Evolution’. (Ed. R. Nielsen.) (Springer: New York.)

Hunter, R. K., Pround, D. N., Burns, J. A., Tibbetts, J. A., and Townsend, V. R. (2007). Parental care in the Neotropical harvestman Phareicranaus calcariferus (Opiliones, Cranaidae). The Journal of Arachnology 35, 199–201.
CrossRef |

Kocher, T. D., Thoma, W. K., Meyer, A., Edwards, S. V., Pääbo, S., Villablanca, F. X., and Wilson, A. C. (1989). Dynamics of mitochondrial DNA evolution in animals: amplification and sequencing with conserved primers. Proceedings of the National Academy of Sciences of the United States of America 86, 6196–6200.
CrossRef |

Kury, A. B. (1993). Análise filogenética de Gonyleptoidea (Arachnida, Opiliones, Laniatores). Ph.D. Thesis, Universidade de São Paulo: São Paulo, Brazil.

Kury, A. B. (2003). Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología Volumen Especial Monográfico 1, 1–337.

Kury, A. B. (2006). Checklist of valid genera of Opiliones of the World, Museu Nacional/UFRJ. Available from http://www.museunacional.ufrj.br/mndi/Aracnologia/checklaniator.htm/ [Accessed 1 February 2011]

Kury, A. B. (2007). Podoctidae Roewer, 1912. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 221–224. (Harvard University Press: Cambridge, MA, USA.)

Kury, A. B., and Cokendolpher, J. C. (2000). Opiliones. In ‘Biodiversidad, Taxonomía y Biogeografía de Artópodos de México: Hacia una Sintesis de su Conocimiento’ vol. 2. (Eds J. E. Llorente, E. González and N. Papavero.) pp. 137–157. (Universidad Nacional Autónoma de Mexico, México DF, Mexico.)

Kury, A. B., and Machado, G. (2009). Notes on Santobius from Vanuatu and Fiji and the status of the eastern Melanesian Ibalonius (Arachnida: Opiliones: Podoctidae). Zoological Studies 48, 524–538.

Kury, A. B., and Pérez G., A. (2002). A new family of Laniatores from northwestern South America (Arachnida, Opiliones). Revista Ibérica de Aracnología 6, 3–11.

Labandeira, C. C., and Sepkoski, J. J. (1993). Insect diversity in the fossil record. Science 261, 310–315.
CrossRef |

Lanyon, S. M. (1988). The stochastic mode of molecular evolution: what consequences for systematic investigations? The Auk 105, 565–573.

Lian, W.-G., Zhu, M.-S., and Kury, A. B. (2008). A new species of the genus Tithaeus from China (Arachnida: Laniatores: Epedanidae). Zootaxa 1841, 53–60.

Machado, G., Requena, G. S., Buzatto, B. A., Osses, F., and Rossetto, L. M. (2004). Five new cases of paternal care in harvestmen (Arachnida: Opiliones): implications for the evolution of male guarding in the Neotropical family Gonyleptidae. Sociobiology 44, 577–598.

Martens, J. (1976). Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida, Opiliones). Entomologica Germanica, Zeitschrift für das Gesamtgebiet der Wissenschaftlichen Entomologie 3, 51–68.

Martens, J. (1980). Versuch eines phylogenetischen Systems der Opiliones. In ‘Proceedings 8th International Congress of Arachnology, Vienna. Verhandlungen’. (Ed. J. Gruber.) pp. 355–360. (Vienna, Austria.)

Martens, J. (1986). Die Grossgliederung der Opiliones und die Evolution der Ordnung (Arachnida). Actas del X Congreso Internacional de Aracnología, Jaca 1, 289–310.

Martens, J. (1993). Further cases of paternal care in Opiliones (Arachnida). Tropical Zoology 6, 97–107.

Martens, J., and Schwendinger, P. S. (1998). A taxonomic revision of the family Oncopodidae I. New genera and new species of Gnomulus Thorell (Opiliones: Laniatores). Revue Suisse de Zoologie 105, 499–555.

Martens, J., Hoheisel, U., and Götze, M. (1981). Vergleichende Anatomie der Legeröhren der Opiliones als Beitrag zur Phylogenie der Ordnung (Arachnida). Zoologische Jahrbucher. Abteilung fur Anatomie und Ontogenie der Tiere 105, 13–76.

Mello-Leitão, C. F. (1949). Famílias, subfamília, espécies e gêneros novos de opiliões e notas de sinonímia. Boletim do Museu Nacional Nova Série, Zoologia, Rio de Janeiro 94, 1–33.

Mendes, A. C., and Kury, A. B. (2007). Stygnopsidae Sørensen, 1932. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 232–234. (Harvard University Press: Cambridge, MA, USA.)

Mendes, A. C., and Kury, A. B. (2008). Intercontinental Triaenonychidae – the case of Ceratomontia (Opiliones, Insidiatores). The Journal of Arachnology 36, 273–279.
CrossRef |

Murphree, C. S. (1988). Morphology of the dorsal integument of ten opilionid species (Arachnida, Opiliones). The Journal of Arachnology 16, 237–252.

Park, J. K., and Ó Foighil, D. (2000). Sphaeriid and corbiculid clams represent separate heterodont bivalve radiations into freshwater environments. Molecular Phylogenetics and Evolution 14, 75–88.
CrossRef |

Pérez González, A. (2006). ‘Revisão Sistemática e Análise Filogenética de Stygnommatidae (Arachnida: Opiliones: Laniatores).’ (Museu Nacional: Universidade Federal de Rio de Janeiro, Rio de Janeiro, Brazil.)

Pérez González, A. (2007). Stygnommatidae Sørensen, 1886. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 229–232. (Harvard University Press: Cambridge, MA, USA.)

Pérez González, A. (2011). New familial assignment for two harvestman species of the infraorder Grassatores (Arachnida: Opiliones: Laniatores). Zootaxa 2757, 24–28.

Pérez González, A., and Kury, A. B. (2007). Samoidae Sørensen, 1886. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 224–226. (Harvard University Press: Cambridge, MA, USA.)

Pineau, P., Henry, M., Suspène, R., Marchio, A., Dettai, A., Debruyne, R., Petit, T., Lécu, A., Moisson, P., Dejean, A., Wain-Hobson, S., and Vartanian, J.-P. (2005). A universal primer set for PCR amplification of nuclear histone H4 genes from all animal species. Molecular Biology and Evolution 22, 582–588.
CrossRef |

Pinto-da-Rocha, R. (1997). Systematic review of the Neotropical family Stygnidae (Opiliones, Laniatores, Gonyleptoidea). Arquivos de Zoologia 33, 163–342.

Pinto-da-Rocha, R., and Kury, A. B. (2003a). Phylogenetic analysis of Santinezia with description of five new species (Opiliones, Laniatores, Cranaidae). The Journal of Arachnology 31, 173–208.
CrossRef |

Pinto-da-Rocha, R., and Kury, A. B. (2003b). Third species of Guasiniidae (Opiliones: Laniatores) with comments on familial relationships. The Journal of Arachnology 31, 394–399.
CrossRef |

Pinto-da-Rocha, R., and Kury, A. B. (2007). Agoristenidae Silhavy, 1973. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 171–173. (Harvard University Press: Cambridge, MA, USA.)

Posada, D. (2005). Modeltest 3.7. Program and documentation available at http://darwin.uvigo.es/ [Accessed 28 May 2011]

Posada, D., and Buckley, T. (2004). Model selection and model averaging in phylogenetics: advantages of Akaike information criterion and Bayesian approaches over likelihood ratio tests. Systematic Biology 53, 793–808.
CrossRef |

Posada, D., and Crandall, K. A. (1998). Modeltest: testing the model of DNA substitution. Bioinformatics (Oxford, England) 14, 817–818.
CrossRef |

Prendini, L., Weygoldt, P., and Wheeler, W. C. (2005). Systematics of the Damon variegates group of African whip spiders (Chelicerata: Amblypygi): evidence from behaviour, morphology and DNA. Organisms, Diversity & Evolution 5, 203–236.
CrossRef |

Rambaut, A. E. (1996). Se-Al sequence alignment editor. University of Oxford, UK Program and documentation available at http://tree.bio.ed.ac.uk/software/figtree/ [Accessed 21 December 2009]

Rambaut, A., and Drummond, A. J. (2009). Tracer v. 1.5. Program and documentation available at http://beast.bio.ed.ac.uk/Tracer [Verified August 2011]

Ree, R. H., and Smith, S. A. (2008). Maximum likelihood inference of geographic range evolution by dispersal, local extinction, and cladogenesis. Systematic Biology 57, 4–14.
CrossRef |

Ree, R. H., Moore, B. R., Webb, C. O., and Donoghue, M. J. (2005). A likelihood framework for inferring the evolution of geographic range on phylogenetic trees. Evolution 59, 2299–2311.

Roewer, C. F. (1912). Die Familien der Assamiiden und Phalangodiden der Opiliones-Laniatores (Assamiden, Dampetriden, Phalangodiden, Epedaniden, Biantiden, Zalmoxiden, Samoiden, Palpidediden anderer Autoren). Archiv für Naturgeschichte, Berlin. Abteilung A 78, 1–242.

Roewer, C. F. (1923). Die Weberknechte der Erde. In ‘Systematische Bearbeitung der bisher bekannten Opiliones’. (Ed. G. Fischer.) pp. 64–120. (Gustav Fischer: Jena, Germany.)

Roewer, C. F. (1949). Über Phalangodiden I: Subfam. Phalangodinae, Tricommatinae, Samoinae; weitere Weberknechte XIII. Senckenbergiana 30, 11–61.

Rokas, A., and Carroll, S. B. (2006). Bushes in the tree of life. PLOS Biology 4, 1899–1904.

Sahney, S., and Benton, M. J. (2008). Recovery from the most profound mass extinction of all time. Proceedings of the Royal Society, Series B 275, 759–765.
CrossRef |

Sanmartín, I., and Ronquist, F. (2004). Southern hemisphere biogeography inferred by event-based models: plant versus animal patterns. Systematic Biology 53, 216–243.
CrossRef |

Schönhofer, A. L., and Martens, J. (2008). Revision of the genus Trogulus Latreille: the Trogulus coriziformis species-group of the western Mediterranean (Opiliones: Trogulidae). Invertebrate Systematics 22, 523–554.
CrossRef |

Schwendinger, P. J., and Giribet, G. (2005). The systematics of the south-east Asian genus Fangensis Rambla (Opiliones: Cyphophthalmi: Stylocellidae). Invertebrate Systematics 19, 297–323.
CrossRef |

Sharma, P., and Giribet, G. (2009). Sandokanid phylogeny based on eight molecular markers – the evolution of a southeast Asian endemic family of Laniatores (Arachnida, Opiliones). Molecular Phylogenetics and Evolution 52, 432–447.
CrossRef |

Sharma, P. P., Prieto, C. E., and Giribet, G. (2011). A new family of Laniatores from the Afrotropics (Arachnida: Opiliones). Invertebrate Systematics 25, 143–154.
CrossRef |

Shultz, J. W., and Regier, J. C. (2001). Phylogenetic analysis of Phalangida (Arachnida, Opiliones) using two nuclear protein-encoding genes supports monophyly of Palpatores. The Journal of Arachnology 29, 189–200.
CrossRef |

Stamatakis, A. (2006). RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics (Oxford, England) 22, 2688–2690.
CrossRef |

Stamatakis, A., Hoover, P., and Rougemont, J. (2008). A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57, 758–771.
CrossRef |

Staręga, W. (1989). Harvestmen (Opiliones) from the Mascarene Islands and the resurrection of the family Zalmoxidae. Annals of the Natal Museum 30, 1–8.

Staręga, W. (1992). An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33, 271–336.

Suzuki, S. (1964). A remarkable new phalangodid, Dongmoa oshimensis, from Japan. Annotationes Zoologicae Japonenses 37, 163–167.

Suzuki, S. (1969). On a collection of Opilionids from Southeast Asia. Journal of Science of the Hiroshima University B 22, 11–77.

Suzuki, S. (1976). Report on a collection of Opilionids from Pasoh Forest Reserve, west Malaysia. Nature and Life in Southeast Asia 7, 9–38.

Suzuki, S. (1977). Report on a collection of opilionids from the Philippines. Journal of Science of the Hiroshima University B 27, 1–120.

Suzuki, S. (1985). A synopsis of the Opiliones of Thailand (Arachnida) I. Cyphopthalmi and Laniatores. Steenstrupia (Copenhagen) 11, 69–110.

Tavaré, S. (1986). Some probabilistic and statistical problems in the analysis of DNA sequences. Lectures on Mathematics in the Life Sciences 17, 57–86.

Thomas, S. M., and Hedin, M. (2008). Multigenic phylogeographic divergence in the paleoendemic southern Appalachian opilionid Fumontana deprehendor Shear (Opiliones, Laniatores, Triaenonychidae). Molecular Phylogenetics and Evolution 46, 645–658.
CrossRef |

Townsend, V. R., Rana, N. J., Proud, D. N., Moore, M. K., Rock, P., and Felgenhauer, B. E. (2009). Morphological changes during postembryonic development in two species of Neotropical harvestmen (Opiliones, Laniatores, Cranaidae). Journal of Morphology 270, 1055–1068.
CrossRef |

Ubick, D., and Dunlop, J. (2005). On the placement of the Baltic amber harvestman Gonyleptes nemastomoides Koch and Berendt, 1854, with notes on the phylogeny of Cladonychiidae (Opiliones, Laniatores, Trvunioidea). Mitteilungen aus dem Museum für Naturkunde in Berlin – Geowissenschaftliche Reihe 8, 75–82.
CrossRef |

Whitfield, J. B., and Lockhart, P. J. (2007). Deciphering ancient rapid radiations. Trends in Ecology and Evolution 22, 258–265.

Whiting, M. F., Carpenter, J. M., Wheeler, Q. D., and Wheeler, W. C. (1997). The Strepsiptera problem: phylogeny of the holometabolous insect orders inferred from 18S and 28S ribosomal DNA sequences and morphology. Systematic Biology 46, 1–68.

Willemart, R. H., Osses, F., Chelini, M. C., Macias-Ordonez, R., and Machado, G. (2009). Sexually dimorphic legs in a Neotropical harvestman (Arachnida, Opiliones): ornament or weapon? Behavioural Processes 80, 51–59.
CrossRef |

Willemart, R. H., Pérez González, A., Farine, J. P., and Gnaspini, P. (2010). Sexually dimorphic tegumental gland openings in Laniatores (Arachnida, Opiliones), with new data on 23 species. Journal of Morphology 271, 641–653.

Xiong, B., and Kocher, T. D. (1991). Comparison of mitochondrial DNA sequences of seven morphospecies of black flies (Diptera: Simuliidae). Genome 34, 306–311.
CrossRef |

Yamaguti, H. Y., and Pinto-da-Rocha, R. (2009). Taxonomic review of Bourguyiinae, cladistic analysis, and a new hypothesis of biogeographic relationships of the Brazilian Atlantic Rainforest (Arachnida: Opiliones, Gonyleptidae). Zoological Journal of the Linnean Society 156, 319–362.
CrossRef |

Yang, Z. (1996). Among-site rate variation and its impact on phylogenetic analyses. Trends in Ecology & Evolution 11, 367–372.
CrossRef |

Zatz, C., Werneck, R. M., Macías-Ordóñez, R., and Machado, G. (2011). Alternative mating tactics in dimorphic males of the harvestman Longiperna concolor (Arachnida: Opiliones). Behavioral Ecology and Sociobiology 65, 995–1005.
CrossRef |

Legal & Privacy | Contact Us | Help


© CSIRO 1996-2016