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Systematics, phylogeny and biogeography
RESEARCH ARTICLE

Phylogeny, evolution and systematic revision of the mite harvestman family Neogoveidae (Opiliones Cyphophthalmi)

Ligia R. Benavides https://orcid.org/0000-0003-4387-1220 A C , Gustavo Hormiga B and Gonzalo Giribet https://orcid.org/0000-0002-5467-8429 A
+ Author Affiliations
- Author Affiliations

A Museum of Comparative Zoology, Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.

B Department of Biological Sciences, The George Washington University, Washington, DC 20052, USA.

C Corresponding author. Email: lbenavidessilva@g.harvard.edu

Invertebrate Systematics 33(1) 101-180 https://doi.org/10.1071/IS18018
Submitted: 6 March 2018  Accepted: 5 August 2018   Published: 6 February 2019

Abstract

Mite harvestmen of the family Neogoveidae have a tropical trans-Atlantic distribution with representatives in equatorial West Africa and the Neotropics, specifically in the south-east region of the USA and in northern South America, being conspicuously absent from Central America. We provide a comprehensive molecular phylogeny of the family including representatives of all genera but the monotypic Tucanogovea Karaman, 2013, and new information on the type species described by Jochen Martens in 1969 that were unavailable for molecular study until now: Brasiliogovea microphaga, Metagovea oviformis and ‘? Gen. enigmaticus’. Additionally, we revisit the somatic and male genitalic morphology of representatives of all genera by means of scanning electron microscopy and confocal laser scanning microscopy, and describe the new genera Leggogovia Benavides & Giribet, gen. nov., Microgovia Benavides, Hormiga & Giribet, gen. nov., Waiwaigovia Benavides, Hormiga & Giribet, gen. nov. and 13 new species: Brasiliogovea aphantostylus Benavides, Hormiga & Giribet, sp. nov., Brasiliogovea microstylus Benavides, Hormiga & Giribet, sp. nov., Brasiliogovea yacambuensis Benavides, Hormiga & Giribet, sp. nov., Metagovea matapi Benavides, Hormiga & Giribet, sp. nov., Metagovea planada Benavides, Hormiga & Giribet, sp. nov., Microgovia chenepau Benavides, Hormiga & Giribet, sp. nov., Neogovea branstetteri Benavides, Hormiga & Giribet, sp. nov., Neogovea enigmatica Martens, sp. nov., Neogovea matawai Benavides, Hormiga & Giribet, sp. nov., Parogovia montealensis Benavides & Giribet, sp. nov., Parogovia prietoi Benavides & Giribet, sp. nov., Parogovia putnami Benavides & Giribet, sp. nov. and Waiwaigovia schultzi Benavides, Hormiga & Giribet, sp. nov. Phylogenetic analyses based on maximum likelihood, parsimony and Bayesian inference support the monophyly of Neogoveidae and a sister group relationship of Neogoveidae + Ogoveidae with Troglosironidae (a clade named Sternophthalmi). Relationships among neogoveid genera are largely congruent between methods as follows: ((Leggogovia gen. nov., Metasiro), (Parogovia, ((Canga, Microgovia gen. nov.), ((Brasiliogovea, Neogovea), (Huitaca, (Waiwaigovia gen. nov., Metagovea)))))). In light of our results, the following taxonomic changes are proposed: Metagovea oviformis Martens, 1969 is transferred to Microgovia, gen. nov.; Parogovia pabsgarnoni Legg, 1990 is transferred to Leggogovia, gen. nov.; ‘? Gen. enigmaticus Martens, 1969’ is an invalid name according to the ICZN; the corresponding taxon is redescribed and formally named as Neogovea enigmatica Martens, sp. nov.

Additional keywords: Afrotropics, arachnids, cryptic fauna, diversity, leaf-litter, Neotropics.


References

Benavides, L. R., and Giribet, G. (2007). An illustrated catalogue of the South American species of the cyphophthalmid family Neogoveidae (Arthropoda, Opiliones, Cyphophthalmi) with a report on 37 undescribed species. Zootaxa 1509, 1–15.

Benavides, L. R., and Giribet, G. (2013). A revision of selected clades of Neotropical mite harvestmen (Arachnida, Opiliones, Cyphophthalmi, Neogoveidae) with the description of eight new species. Bulletin of the Museum of Comparative Zoology 161, 1–44.
A revision of selected clades of Neotropical mite harvestmen (Arachnida, Opiliones, Cyphophthalmi, Neogoveidae) with the description of eight new species.Crossref | GoogleScholarGoogle Scholar |

Boyer, S. L., Clouse, R. M., Benavides, L. R., Sharma, P., Schwendinger, P. J., Karunarathna, I., and Giribet, G. (2007). Biogeography of the world: a case study from cyphophthalmid Opiliones, a globally distributed group of arachnids. Journal of Biogeography 34, 2070–2085.
Biogeography of the world: a case study from cyphophthalmid Opiliones, a globally distributed group of arachnids.Crossref | GoogleScholarGoogle Scholar |

Clouse, R. M., and Wheeler, W. C. (2014). Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates. Zootaxa 3814, 177–201.
Descriptions of two new, cryptic species of Metasiro (Arachnida: Opiliones: Cyphophthalmi: Neogoveidae) from South Carolina, USA, including a discussion of mitochondrial mutation rates.Crossref | GoogleScholarGoogle Scholar |

Clouse, R. M., Sharma, P. P., Stuart, J. C., Davis, L. R., Giribet, G., Boyer, S. L., and Wheeler, W. C. (2016). Phylogeography of the harvestman genus Metasiro (Arthropoda, Arachnida, Opiliones) reveals a potential solution to the Pangean paradox. Organisms, Diversity & Evolution 16, 167–184.
Phylogeography of the harvestman genus Metasiro (Arthropoda, Arachnida, Opiliones) reveals a potential solution to the Pangean paradox.Crossref | GoogleScholarGoogle Scholar |

DaSilva, M. B., Pinto-da-Rocha, R., and Giribet, G. (2010). Canga renatae, a new genus and species of Cyphophthalmi from Brazilian Amazon caves (Opiliones: Neogoveidae). Zootaxa 2508, 45–55.

Davis, N. W. (1933). A new opilionid from Florida (Arachnida, Cyphophthalmi). Journal of the New York Entomological Society 41, 49–53.

Davis, N. W. (1937). A cyphophthalmid from South America. (Arachnida, Phalangida). Journal of the New York Entomological Society 45, 133–137.

de Bivort, B. L., and Giribet, G. (2004). A new genus of cyphophthalmid from the Iberian Peninsula with a phylogenetic analysis of the Sironidae (Arachnida: Opiliones: Cyphophthalmi) and a SEM database of external morphology. Invertebrate Systematics 18, 7–52.
A new genus of cyphophthalmid from the Iberian Peninsula with a phylogenetic analysis of the Sironidae (Arachnida: Opiliones: Cyphophthalmi) and a SEM database of external morphology.Crossref | GoogleScholarGoogle Scholar |

de Oliveira Bernardi, L. F., Klompen, H., Zacarias, M. S., and Ferreira, R. L. (2013). A new species of Neocarus Chamberlin & Mulaik, 1942 (Opilioacarida, Opilioacaridae) from Brazil, with remarks on its postlarval development. ZooKeys 358, 69–89.
A new species of Neocarus Chamberlin & Mulaik, 1942 (Opilioacarida, Opilioacaridae) from Brazil, with remarks on its postlarval development.Crossref | GoogleScholarGoogle Scholar |

Farris, J. S., Albert, V. A., Källersjö, M., Lipscomb, D., and Kluge, A. G. (1996). Parsimony jackknifing outperforms neighbor-joining. Cladistics 12, 99–124.
Parsimony jackknifing outperforms neighbor-joining.Crossref | GoogleScholarGoogle Scholar |

Fernández, R., and Giribet, G. (2015). Unnoticed in the tropics: phylogenomic resolution of the poorly known arachnid order Ricinulei (Arachnida). Royal Society Open Science 2, 150065.
Unnoticed in the tropics: phylogenomic resolution of the poorly known arachnid order Ricinulei (Arachnida).Crossref | GoogleScholarGoogle Scholar | 26543583PubMed |

Giribet, G. (2000). Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). Revista Ibérica de Aracnología 2, 49–76.

Giribet, G. (2003). Stability in phylogenetic formulations and its relationship to nodal support. Systematic Biology 52, 554–564.
Stability in phylogenetic formulations and its relationship to nodal support.Crossref | GoogleScholarGoogle Scholar | 12857646PubMed |

Giribet, G. (2007). Efficient tree searches with available algorithms. Evolutionary Bioinformatics Online 3, 341–356.
Efficient tree searches with available algorithms.Crossref | GoogleScholarGoogle Scholar | 19461977PubMed |

Giribet, G., and Prieto, C. E. (2003). A new Afrotropical Ogovea (Opiliones, Cyphophthalmi) from Cameroon, with a discussion on the taxonomic characters in the family Ogoveidae. Zootaxa 329, 1–18.

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.
A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores.Crossref | GoogleScholarGoogle Scholar |

Giribet, G., Sharma, P. P., Benavides, L. R., Boyer, S. L., Clouse, R. M., de Bivort, B. L., Dimitrov, D., Kawauchi, G. Y., Murienne, J. Y., and Schwendinger, P. J. (2012). Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement. Biological Journal of the Linnean Society. Linnean Society of London 105, 92–130.
Evolutionary and biogeographical history of an ancient and global group of arachnids (Arachnida: Opiliones: Cyphophthalmi) with a new taxonomic arrangement.Crossref | GoogleScholarGoogle Scholar |

Giupponi, A. P. L., and Kury, A. B. (2015). A new species of Metagovea Rosas Costa, 1950 from Napo Province, Ecuador (Opiliones, Cyphophthalmi, Neogoveidae). ZooKeys 477, 1–15.
A new species of Metagovea Rosas Costa, 1950 from Napo Province, Ecuador (Opiliones, Cyphophthalmi, Neogoveidae).Crossref | GoogleScholarGoogle Scholar |

Goloboff, P. A. (1999). Analyzing large data sets in reasonable times: solutions for composite optima. Cladistics 15, 415–428.
Analyzing large data sets in reasonable times: solutions for composite optima.Crossref | GoogleScholarGoogle Scholar |

Goloboff, P. A. (2002). Techniques for analyzing large data sets. In ‘Techniques in Molecular Systematics and Evolution’. (Eds R. DeSalle, G. Giribet and W. Wheeler.) pp. 70–79. (Brikhäuser Verlag: Basel, Switzerland.)

Goodnight, C. J., and Goodnight, M. L. (1942). Phalangids from British Guiana. American Museum Novitates 1167, 1–13.

Goodnight, C. J., and Goodnight, M. L. (1980). Metagovea philipi, n. sp., a new cyphophthalmid (Arachnida) from Ecuador. Transactions of the American Microscopical Society 99, 128–131.
Metagovea philipi, n. sp., a new cyphophthalmid (Arachnida) from Ecuador.Crossref | GoogleScholarGoogle Scholar |

Hall, T. A. (1999). BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series 41, 95–98.

Hansen, H. J. (1921). ‘Studies on Arthropoda I. The Pedipalpi, Ricinulei, and Opiliones (exc. Op. Laniatores) collected by Mr. Leonardo Fea in tropical West Africa and adjacent Islands.’ (Gyldendalske Boghandel: Copenhagen, Denmark.)

Hinton, H. E. (1938). A key to the genera of the suborder Cyphophthalmi with a description and figures of Neogovea immsi, gen. et sp.n. (Arachnida, Opiliones). Annals and Magazine of Natural History, Ser.11 2, 331–338.
A key to the genera of the suborder Cyphophthalmi with a description and figures of Neogovea immsi, gen. et sp.n. (Arachnida, Opiliones).Crossref | GoogleScholarGoogle Scholar |

Hiřman, M., Kotyk, M., Kotyková Varadínová,, Z., and Šťáhlavský, (2018). First cytogenetic study of a member of the harvestman family Neogoveidae (Opiliones: Cyphophthalmi: Sternophthalmi) from Cameroon with description of a new species Parogovia parasironoides sp. nov. Annales Zoologici 68, 867–877.

Hoffman, R. L. (1963). A new phalangid of the genus Siro from eastern United States, and taxonomic notes on other American sironids (Arach., Opiliones). Senckenbergiana Biologica 44, 129–139.

Jocqué, M., and Jocqué, R. (2011). An overview of Neogovea species (Opiliones: Cyphophthalmi: Neogoveidae) with the description of Neogovea virginie n. sp. from French Guiana. Zootaxa 2754, 41–50.

Juberthie, C. (1960). Contribution a l’étude des opilions cyphophthalmes: description de Metasiro gen. n. Bulletin du Muséum National d’Histoire Naturelle, 2e série 32, 235–241.

Juberthie, C. (1969). Sur les opilions cyphophthalmes Stylocellinae du Gabon. Biologia Gabonica 5, 79–92.

Juberthie, C. (1970). Les genres d’opilions Sironinae (Cyphophthalmes). Bulletin du Muséum National d’Histoire Naturelle, 2e Série 41, 1371–1390.

Karaman, I. (2013). Tucanogovea schusteri n. gen. n. sp., a new cyphophthalmid (Opiliones, Cyphophthalmi, Neogoveidae) from Amazonia. Biologia Serbica 35, 68–75.

Katoh, K., and Standley, D. M. (2013). MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Molecular Biology and Evolution 30, 772–780.
MAFFT multiple sequence alignment software version 7: improvements in performance and usability.Crossref | GoogleScholarGoogle Scholar | 23329690PubMed |

Lanfear, R., Calcott, B., Ho, S. Y., and Guindon, S. (2012). PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29, 1695–1701.
PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses.Crossref | GoogleScholarGoogle Scholar | 22319168PubMed |

Legg, G. (1990). Parogovia pabsgarnoni, sp. n. (Arachnida, Opiliones, Cyphophthalmi) from Sierra Leone, with notes on other African species of Parogovia. Bulletin of the British Arachnological Society 8, 113–121.

Machado, G., and Macías-Ordóñez, R. (2007). Reproduction. In ‘Harvestmen: the Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 414–454. (Harvard University Press: Cambridge, MA.)

Martens, J. (1969). Cyphophthalmi aus Brasilien (Opiliones). Beiträge zur Neotropischen Fauna 6, 109–119.
Cyphophthalmi aus Brasilien (Opiliones).Crossref | GoogleScholarGoogle Scholar |

Mickevich, M. F., and Farris, J. S. (1981). The implications of congruence in Menidia. Systematic Zoology 30, 351–370.
The implications of congruence in Menidia.Crossref | GoogleScholarGoogle Scholar |

Miller, M. A., Pfeiffer, W., and Schwartz, T. (2010). Creating the CIPRES science gateway for inference of large phylogenetic trees. In ‘Proceedings of the Gateway Computing Environments Workshop (GCE)’. New Orleans pp. 1–8.

Murienne, J., Benavides, L. R., Prendini, L., Hormiga, G., and Giribet, G. (2013). Forest refugia in western and central Africa as ‘museums’ of Mesozoic biodiversity. Biology Letters 9, 20120932.
Forest refugia in western and central Africa as ‘museums’ of Mesozoic biodiversity.Crossref | GoogleScholarGoogle Scholar | 23193047PubMed |

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

Nixon, K. C. (1999). The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics 15, 407–414.
The parsimony ratchet, a new method for rapid parsimony analysis.Crossref | GoogleScholarGoogle Scholar |

Owen-Smith, T. M., Ganerød, M., van Hinsbergen, D. J. J., Gaina, C., Ashwal, L. D., and Torsvik, T. H. (2017). Testing Early Cretaceous Africa–South America fits with new palaeomagnetic data from the Etendeka Magmatic Province (Namibia). Tectonophysics , .
Testing Early Cretaceous Africa–South America fits with new palaeomagnetic data from the Etendeka Magmatic Province (Namibia).Crossref | GoogleScholarGoogle Scholar |

Pinto-da-Rocha, R., and Andrade, R. (2012). A new species of Cryptocellus (Arachnida: Ricinulei) from eastern Amazonia. Zoologia (Curitiba) 29, 474–478.
A new species of Cryptocellus (Arachnida: Ricinulei) from eastern Amazonia.Crossref | GoogleScholarGoogle Scholar |

Rambaut, A., Suchard, M. A., Xie, D., and Drummond, A. J. (2003–2014). Tracer: MCMC Trace Analysis Tool Version 1.6.1pre, Available from http://beast.bio.ed.ac.uk/Tracer [verified 19 December 2018].

Rambla, M. and Juberthie, C. (1994). Opiliones. In ‘Encyclopaedia Biospeologica’. (Eds C. Juberthie and V. Decu.) pp. 215–230. (Société de Biospéologie: Moulis-Boucarest, France.)

Roewer, C. F. (1927). Weitere Weberknechte I. Ergänzung der: ‘Webernechte der Erde’, 1923. Abhandlungen Naturwissenschaftlichen Verein zu Bremen 26, 261–402.

Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D. L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M. A., and Huelsenbeck, J. P. (2012). MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61, 539–542.
MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space.Crossref | GoogleScholarGoogle Scholar | 22357727PubMed |

Rosas Costa, J. A. (1950). Sinopsis de los géneros de Sironinae, con la descripción de dos géneros y una especie nuevos (Opiliones, Cyphophthalmi). Arthropoda 1, 127–151.

Sharma, P., and Giribet, G. (2009). A relict in New Caledonia: phylogenetic relationships of the family Troglosironidae (Opiliones: Cyphophthalmi). Cladistics 25, 279–294.
A relict in New Caledonia: phylogenetic relationships of the family Troglosironidae (Opiliones: Cyphophthalmi).Crossref | GoogleScholarGoogle Scholar |

Sharma, P. P., Vahtera, V., Kawauchi, G. Y., and Giribet, G. (2011). Running WILD: the case for exploring mixed parameter sets in sensitivity analysis. Cladistics 27, 538–549.
Running WILD: the case for exploring mixed parameter sets in sensitivity analysis.Crossref | GoogleScholarGoogle Scholar |

Shear, W. A. (1977). The opilionid genus Neogovea Hinton, with a description of the first troglobitic cyphophthalmid from the western hemisphere (Opiliones, Cyphophthalmi). The Journal of Arachnology 3, 165–175.

Shear, W. A. (1979). Huitaca ventralis, n. gen., n. sp., with a description of a gland complex new to cyphophthalmids (Opiliones, Cyphophthalmi). The Journal of Arachnology 7, 237–243.

Shear, W.A. (1980). A review of the Cyphophthalmi of the United States and Mexico, with a proposed reclassification of the suborder (Arachnida, Opiliones). American Museum Novitates 2705, 1–34.

Souza, M. F. V. R., and Lopes Ferreira, R. (2010). Eukoenenia (Palpigradi: Eukoeneniidae) in Brazilian caves with the first troglobiotic palpigrade from South America. The Journal of Arachnology 38, 415–424.
Eukoenenia (Palpigradi: Eukoeneniidae) in Brazilian caves with the first troglobiotic palpigrade from South America.Crossref | GoogleScholarGoogle Scholar |

Stamatakis, A. (2014). RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30, 1312–1313.
RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies.Crossref | GoogleScholarGoogle Scholar | 24451623PubMed |

Stamatakis, A., Hoover, P., and Rougemont, J. (2008). A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57, 758–771.
A rapid bootstrap algorithm for the RAxML web servers.Crossref | GoogleScholarGoogle Scholar | 18853362PubMed |

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

Svensen, H. H., Torsvik, T. H., Callegaro, S., Augland, L., Heimdal, T. H., Jerram, D. A., Planke, S., and Pereira, E. (2017). Gondwana large igneous provinces: plate reconstructions, volcanic basins and sill volumes. pp. SP463.7. (The Geological Society of London: London.)

Torsvik, T. H., Rousse, S., Labails, C., and Smethurst, M. A. (2009). A new scheme for the opening of the South Atlantic Ocean and the dissection of an Aptian salt basin. Geophysical Journal International 177, 1315–1333.
A new scheme for the opening of the South Atlantic Ocean and the dissection of an Aptian salt basin.Crossref | GoogleScholarGoogle Scholar |

Tsurusaki, N. (2007). Cytogenetics. In ‘Harvestmen: the Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 266–279. (Harvard University Press: Cambridge, MA.)

Vaidya, G., Lohman, D. J., and Meier, R. (2011). SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information. Cladistics 27, 171–180.
SequenceMatrix: concatenation software for the fast assembly of multi-gene datasets with character set and codon information.Crossref | GoogleScholarGoogle Scholar |

Wheeler, W. C. (1995). Sequence alignment, parameter sensitivity, and the phylogenetic analysis of molecular data. Systematic Biology 44, 321–331.
Sequence alignment, parameter sensitivity, and the phylogenetic analysis of molecular data.Crossref | GoogleScholarGoogle Scholar |

Wheeler, W. C. (1999). Measuring topological congruence by extending character techniques. Cladistics 15, 131–135.
Measuring topological congruence by extending character techniques.Crossref | GoogleScholarGoogle Scholar |

Wheeler, W. C., Lucaroni, N., Hong, L., Crowley, L. M., and Varón, A. (2015). POY version 5: phylogenetic analysis using dynamic homologies under multiple optimality criteria. Cladistics 31, 189–196.
POY version 5: phylogenetic analysis using dynamic homologies under multiple optimality criteria.Crossref | GoogleScholarGoogle Scholar |

Willemart, R. H., and Giribet, G. (2010). A scanning electron microscopic survey of the cuticle in Cyphophthalmi (Arachnida, Opiliones) with the description of novel sensory and glandular structures. Zoomorphology 129, 175–183.
A scanning electron microscopic survey of the cuticle in Cyphophthalmi (Arachnida, Opiliones) with the description of novel sensory and glandular structures.Crossref | GoogleScholarGoogle Scholar |

Zhang, J., Kapli, P., Pavlidis, P., and Stamatakis, A. (2013). A general species delimitation method with applications to phylogenetic placements. Bioinformatics 29, 2869–2876.
A general species delimitation method with applications to phylogenetic placements.Crossref | GoogleScholarGoogle Scholar | 23990417PubMed |