A systematic revision of the South African Pettalidae (Arachnida : Opiliones : Cyphophthalmi) based on a combined analysis of discrete and continuous morphological characters with the description of seven new species
Benjamin L. de Bivort A B C and Gonzalo Giribet BA Rowland Institute at Harvard, Harvard University, 100 Edwin Land Boulevard, Cambridge, MA 02142, USA.
B Museum of Comparative Zoology and Department of Organismic and Evolutionary Biology, Harvard University, 26 Oxford Street, Cambridge, MA 02138, USA.
C Corresponding author. Email: debivort@rowland.harvard.edu
Invertebrate Systematics 24(4) 371-406 https://doi.org/10.1071/IS10015
Submitted: 23 May 2010 Accepted: 23 September 2010 Published: 13 December 2010
Abstract
The cyphophthalmid family Pettalidae in South Africa is revised and seven new species are described from museum material collected between 1939 and 1985. Two of these are placed in the genus Purcellia and five in Parapurcellia, bringing the total number of described South African cyphophthalmids to 15. In addition, Purcellia peregrinator is transferred to the genus Parapurcellia. Phylogenetic analyses of discrete morphological and continuous morphometric characters, both separately and in combination, support the generic assignments and contribute towards a more detailed understanding of the systematics of the group in South Africa. In order to assess the stability of our phylogenetic results, the different morphological datasets were analysed under equal and implied weighting, as well as under several weighting schemes that varied the respective contribution to tree length of the discrete and continuous data partitions. These variations generated two phylogenetic hypotheses: (1) monophyly of the South African pettalids + Austropurcellia from north-eastern Australia as a derived clade within Pettalidae; and (2) polyphyly of the South African pettalids with Parapurcellia basal within Pettalidae. The latter hypothesis is congruent with previous molecular phylogenies of Cyphophthalmi, and has moderate bootstrap support. The sisterhood of Purcellia griswoldi, sp. nov. and P. lawrencei, sp. nov. receives high nodal support across analytic methods. New combination: Parapurcellia peregrinator (Lawrence, 1963).
Additional keywords: Afromontane forests, Gondwanan biogeography, morphometrics, South Africa.
References
Boyer, S. L., and Giribet, G. (2007). A new model Gondwanan taxon: systematics and biogeography of the harvestman family Pettalidae (Arachnida, Opiliones, Cyphophthalmi), with a taxonomic revision of genera from Australia and New Zealand. Cladistics 23, 337–361.| A new model Gondwanan taxon: systematics and biogeography of the harvestman family Pettalidae (Arachnida, Opiliones, Cyphophthalmi), with a taxonomic revision of genera from Australia and New Zealand.Crossref | GoogleScholarGoogle Scholar |
Boyer, S. L., and Giribet, G. (2009). Welcome back New Zealand: regional biogeography and Gondwanan origin of three endemic genera of mite harvestmen (Arachnida, Opiliones, Cyphophthalmi). Journal of Biogeography 36, 1084–1099.
| Welcome back New Zealand: regional biogeography and Gondwanan origin of three endemic genera of mite harvestmen (Arachnida, Opiliones, Cyphophthalmi).Crossref | GoogleScholarGoogle Scholar |
Boyer, S. L., Baker, J. M., and Giribet, G. (2007a). Deep genetic divergences in Aoraki denticulata (Arachnida, Opiliones, Cyphophthalmi): a widespread ‘mite harvestman’ defies DNA taxonomy. Molecular Ecology 16, 4999–5016.
| Deep genetic divergences in Aoraki denticulata (Arachnida, Opiliones, Cyphophthalmi): a widespread ‘mite harvestman’ defies DNA taxonomy.Crossref | GoogleScholarGoogle Scholar | 17944852PubMed |
Boyer, S. L., Clouse, R. M., Benavides, L. R., Sharma, P., Schwendinger, P. J., Karunarathna, I., and Giribet, G. (2007b). 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 Giribet, G. (2007). Across Lydekker’s line-first report of mite harvestmen (Opiliones : Cyphophthalmi : Stylocellidae) from New Guinea. Invertebrate Systematics 21, 207–228.
| Across Lydekker’s line-first report of mite harvestmen (Opiliones : Cyphophthalmi : Stylocellidae) from New Guinea.Crossref | GoogleScholarGoogle Scholar |
Clouse, R. M., de Bivort, B. L., and Giribet, G. (2009). A phylogenetic analysis for the south-east Asian mite harvestman family Stylocellidae (Opiliones: Cyphophthalmi) – a combined analysis using morphometric and molecular data. Invertebrate Systematics 23, 515–529.
| A phylogenetic analysis for the south-east Asian mite harvestman family Stylocellidae (Opiliones: Cyphophthalmi) – a combined analysis using morphometric and molecular data.Crossref | GoogleScholarGoogle Scholar |
Daniels, S. R., Picker, M. D., Cowlin, R. M., and Hamer, M. L. (2009). Unravelling evolutionary lineages among South African velvet worms (Onychophora: Peripatopsis) provides evidence for widespread cryptic speciation. Biological Journal of the Linnean Society. Linnean Society of London 97, 200–216.
| Unravelling evolutionary lineages among South African velvet worms (Onychophora: Peripatopsis) provides evidence for widespread cryptic speciation.Crossref | GoogleScholarGoogle Scholar |
de Bivort, B., 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 Bivort, B. L., Clouse, R. M., and Giribet, G. (2010). A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen (Opiliones, Cyphophthalmi, Pettalidae). Journal of Zoological Systematics and Evolutionary Research 48, 294–309.
| A morphometrics-based phylogeny of the temperate Gondwanan mite harvestmen (Opiliones, Cyphophthalmi, Pettalidae).Crossref | GoogleScholarGoogle Scholar |
Ewing, H. E. (1923). Holosiro acaroides, new genus and species, the only new world representative of the mite-like phalangids of the suborder Cyphophthalmi. Annals of the Entomological Society of America 16, 387–391.
Forster, R. R. (1948). The sub-order Cyphophthalmi Simon in New Zealand. Dominion Museum Records in Entomology 1, 79–119.
Forster, R. R. (1952). Supplement to the sub-order Cyphophthalmi. Dominion Museum Records in Entomology 1, 179–211.
Forster, R. R. (1955). Further Australian harvestmen (Arachnida: Opiliones). Australian Journal of Zoology 3, 354–411.
| Further Australian harvestmen (Arachnida: Opiliones).Crossref | GoogleScholarGoogle Scholar |
Giribet, G. (2000). Catalogue of the Cyphophthalmi of the world (Arachnida, Opiliones). Revista Ibérica de Aracnología 2, 49–76.
Giribet, G. (2002). Stylocellus ramblae, a new stylocellid (Opiliones, Cyphophthalmi) from Singapore, with a discussion of the family Stylocellidae. The Journal of Arachnology 30, 1–9.
| Stylocellus ramblae, a new stylocellid (Opiliones, Cyphophthalmi) from Singapore, with a discussion of the family Stylocellidae.Crossref | GoogleScholarGoogle Scholar |
Giribet, G. (2003). Karripurcellia, a new pettalid genus (Arachnida : Opiliones : Cyphophthalmi) from Western Australia, with a cladistic analysis of the family Pettalidae. Invertebrate Systematics 17, 387–406.
| Karripurcellia, a new pettalid genus (Arachnida : Opiliones : Cyphophthalmi) from Western Australia, with a cladistic analysis of the family Pettalidae.Crossref | GoogleScholarGoogle Scholar |
Giribet, G. (2005). Book reviews: TNT: tree analysis using new technology. Systematic Biology 54, 176–178.
| Book reviews: TNT: tree analysis using new technology.Crossref | GoogleScholarGoogle Scholar |
Giribet, G. (2007). Efficient tree searches with available algorithms. Evolutionary Bioinformatics 3, 1–16.
Giribet, G., and Boyer, S. L. (2002). A cladistic analysis of the cyphophthalmid genera (Opiliones, Cyphophthalmi). The Journal of Arachnology 30, 110–128.
| A cladistic analysis of the cyphophthalmid genera (Opiliones, Cyphophthalmi).Crossref | GoogleScholarGoogle Scholar |
Giribet, G., and Boyer, S. L. (2007). Pettalidae Shear, 1980. In ‘Harvestmen: The Biology of Opiliones’. (Eds R. Pinto-da-Rocha, G. Machado and G. Giribet.) pp. 99–101. (Harvard University Press: Cambridge, MA.)
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 |
Goloboff, P. A. (1993). Estimating character weights during tree search. Cladistics 9, 83–91.
| Estimating character weights during tree search.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., Farris, J. S., and Nixon, K. C. (2008). TNT, a free program for phylogenetic analyses. Cladistics 24, 774–786.
| TNT, a free program for phylogenetic analyses.Crossref | GoogleScholarGoogle Scholar |
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 |
Griswold, C. E. (1991). Cladistic biogeography of Afromontane spiders. Australian Systematic Botany 4, 73–89.
| Cladistic biogeography of Afromontane spiders.Crossref | GoogleScholarGoogle Scholar |
Hamer, M. L., Samways, M. J., and Ruhberg, H. (1997). A review of the Onychophora of South Africa, with discussion of their conservation. Annals of the Natal Museum 38, 283–312.
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.)
Hansen, H. J., and Sørensen, W. (1904). ‘On two orders of Arachnida: Especially the Suborder Cyphophthalmi, and Ricinulei, Namely the Family Cryptostemmatoidae.’ (Cambridge University Press: Cambridge, UK.)
Harvey, M. S. (1996). Small arachnids and their value in Gondwanan biogeographic studies. In ‘Gondwanan Heritage: Past, Present and Future of Western Australian Biota’. (Ed. S. D. Hopper.) pp. 155–162. (Surrey Beatty & Sons: Sydney.)
Hirst, S. (1925). On some new genera and species of Arachnida. Proceedings of the Zoological Society of London 1925, 1271–1280.
Joseph, G. (1868). Cyphophthalmus duricorius, eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdeckt in der Luëger Grotte in Krain. Berliner Entomologische Zeitung 12, 241–250.
| Cyphophthalmus duricorius, eine neue Arachniden-Gattung aus einer neuen Familie der Arthrogastren-Ordnung entdeckt in der Luëger Grotte in Krain.Crossref | GoogleScholarGoogle Scholar |
Juberthie, C. (1956). Une nouvelle espèce d’Opilions Sironidae de France et d’Espagne: Parasiro coiffaiti n. sp. Bulletin du Muséum national d’histoire naturelle, Paris 28, 394–400.
Juberthie, C. (1969). Sur les opilions cyphophthalmes Stylocellinae du Gabon. Biologia Gabonica 5, 79–92.
Juberthie, C. (1970). Les opilions cyphophthalmes cavernicoles. Notes sur Speleosiro argasiformis Lawrence. Bulletin du Museum National d’Histoire Naturelle 42, 864–871.
Juberthie, C. (1979). Un cyphophthalme nouveau d’une grotte de Nouvelle-Caledonie: Troglosiro aelleni n. gen., n. sp. (Opilion, Sironinae). Revue Suisse de Zoologie 86, 221–231.
Juberthie, C. (1988). Un nouvel opilion cyphophthalme aveugle d’Australie: Austropurcellia gen. nov., scoparia n. sp. Memoires de Biospeologie 15, 133–140.
Kauri, H. (1961). Opiliones. In ‘South African Animal Life. Results of the Lund University Expedition in 1950–1951’. (Eds B. Hanström, P. Brinck and G. Rudebeck.) pp. 9–197. (Almquist & Wiksell: Uppsala.)
Kury, A. B. (2006). A new species of Graemontia Lawrence, 1931, from the Western Cape, South Africa, with notes on the relationships of the genus (Opiliones: Laniatores: Triaenonychidae). African Zoology 41, 45–50.
| A new species of Graemontia Lawrence, 1931, from the Western Cape, South Africa, with notes on the relationships of the genus (Opiliones: Laniatores: Triaenonychidae).Crossref | GoogleScholarGoogle Scholar |
Latreille, P. A. (1804). ‘Histoire naturelle, generale et particulière des Crustacés et des Insectes.’ (F. Duart: Paris.)
Lawrence, R. F. (1931). The harvest-spiders (Opiliones) of South Africa. Annals of the South African Museum 29, 341–508.
Lawrence, R. F. (1933). The harvest-spiders (Opiliones) of Natal. Annals of the Natal Museum 7, 211–241.
Lawrence, R. F. (1939). A contribution to the opilionid fauna of Natal and Zululand. Annals of the Natal Museum 9, 225–243.
Lawrence, R. F. (1963). The Opiliones of the Transvaal. Annals of the Transvaal Museum 24, 275–304.
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.
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 |
Pickard-Cambridge, O. (1875). On three new and curious forms of Arachnida. Annals and Magazine of Natural History. Series 4, 383–390.
Roewer, C. F. (1916). 7 neue Opilioniden des Zoolog. Museums in Berlin. Archiv für Naturgeschichte 81, 6–13.
Roewer, C. F. (1923). ‘Die Weberknechte der Erde. Systematische Bearbeitung der bisher bekannten Opiliones.’ (Verlag von Gustav Fisher: Jena.)
Roewer, C. F. (1961). Opiliones aus Sud-Chile. Senckenbergiana Biologica 42, 99–105.
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.
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.
| The systematics of the south-east Asian genus Fangensis Rambla (Opiliones : Cyphophthalmi : Stylocellidae).Crossref | GoogleScholarGoogle Scholar |
Sharma, P., and Giribet, G. (2005). A new Troglosiro species (Opiliones, Cyphophthalmi, Troglosironidae) from New Caledonia. Zootaxa 1053, 47–60.
Sharma, P., and Giribet, G. (2006). A new Pettalus species (Opiliones, Cyphophthalmi, Pettalidae) from Sri Lanka with a discussion on the evolution of eyes in Cyphophthalmi. The Journal of Arachnology 34, 331–341.
| A new Pettalus species (Opiliones, Cyphophthalmi, Pettalidae) from Sri Lanka with a discussion on the evolution of eyes in Cyphophthalmi.Crossref | GoogleScholarGoogle Scholar |
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, 237–242.
Shear, W. A. (1985). Marwe coarctata, a remarkable new cyphophthalmid from a limestone cave in Kenya (Arachnida, Opiliones). American Museum Novitates 2830, 1–6.
Shear, W. A. (1993a). The genus Chileogovea (Opiliones, Cyphophthalmi, Pettalidae). The Journal of Arachnology 21, 73–78.
Shear, W. A. (1993b). The genus Troglosiro and the new family Troglosironidae (Opiliones, Cyphophthalmi). The Journal of Arachnology 21, 81–90.
Shear, W. A. (1993c). New species in the opilionid genus Stylocellus from Malaysia, Indonesia and the Philippines (Opiliones, Cyphophthalmi, Stylocellidae). Bulletin of the British Arachnological Society 9, 174–188.
Shear, W. A., and Gruber, J. (1996). Cyphophthalmid opilionids new to Madagascar: two new genera (Opiliones, Cyphophthalmi, Pettalidae). Bulletin of the British Arachnological Society 10, 181–186.
Starega, W. (2008). The second species of Purcellia Hansen et Sørensen, 1904 (Arachnida: Opiliones: Pettalidae) from South Africa. Polish Journal of Entomology 77, 51–56.
Thorell, T. (1876). Sopra alcuni Opilioni (Phalangidea) d’Europa e dell’Asia occidentale, con un quadro dei generi europei de quest’Ordine. Annali Museo civico Storia naturale Giacomo Doria 8, 452–508.