The evolutionary history of Lipsothrix Loew (Diptera : Tipuloidea) inferred through systematic revision and historical biogeographical analysis
Matthew PetersenRoanoke College, Biology Department, 221 College Lane, Salem, VA 24153, USA. Email: mpetersen13@gmail.com
Invertebrate Systematics 29(3) 239-286 https://doi.org/10.1071/IS14044
Submitted: 14 August 2014 Accepted: 15 February 2015 Published: 30 June 2015
Abstract
The evolutionary history of the crane fly genus Lipsothrix Lowe, 1873 was examined through a systematic revision of specimens maintained in natural history collections, parsimony analysis using TNT based on immature and adult morphology, and dispersal–vicariance analyses in RASP (Bayesian and S-DIVA). The genus Lipsothrix was found to contain 29 valid species, one subspecies (L. nobilis iranica) and three species maintained but in need of further evaluation (Lipsothrix burmica, L. heitfeldi, L. yakushimae). One species complex (L. shasta complex) was further resolved using ordination and cluster analyses. One new species is described, L. nullusarma, sp. nov., and three species are synonymised (L. fulva, L. mirifica, L. yamamotoana omogoensis). Phylogenetic analysis recovered a strongly supported and monophyletic Lipsothrix. The historical biogeographical analyses of Lipsothrix inferred an ancestral area in the Western Palearctic region that likely dates to the Eocene. One, or several, dispersal events occurred from the Western Palearctic before major diversification events occurred in the Oriental and Eastern Palearctic + Nearctic regions. Two major paleogeographical developments occurring in the Oriental and Eastern Palearctic + Nearctic regions are largely responsible for Lipsothrix diversification, the collision of the Indian plate with Asia and the rise of the trans-Beringian land bridges during the Tertiary and Quaternary.
References
Alexander, C. P. (1916). New or little-known crane-flies from the United States and Canada: Tipulidae, Ptychopteridae, Diptera. Part 3. Proceedings. Academy of Natural Sciences of Philadelphia 68, 486–549.Alexander, C. P. (1928). New or little-known Tipulidae from eastern Asia (Diptera), II. Philippine Journal of Science 35, 455–489.
Alexander, C. P. (1929). New or little-known Tipulidae from eastern Asia (Diptera), V. Philippine Journal of Science 40, 519–547.
Alexander, C. P. (1930). New or little-known Tipulidae from eastern Asia (Diptera). VI. Philippine Journal of Science 42, 59–83.
Alexander, C. P. (1931). Crane-flies of the Baltic Amber (Diptera). Bernstein-Forschungen 2, 1–135.
Alexander, C. P. (1933). New or little-known Tipulidae from eastern Asia (Diptera), XII. Philippine Journal of Science 50, 129–162.
Alexander, C. P. (1935). New or little-known Tipulidae from eastern Asia (Diptera), XXIII. Philippine Journal of Science 56, 339–372.
Alexander, C. P. (1938). New or little-known Tipulidae from eastern Asia (Diptera), XXXVII. Philippine Journal of Science 66, 221–259.
Alexander, C. P. (1940). New or little-known Tipulidae form eastern Asia (Diptera), XLII. Philippine Journal of Science 71, 169–204.
Alexander, C. P. (1942). Familly Tipulidae. Bulletin of the Connecticut State Geological and Natural History Survey 64, 196–509.
Alexander, C. P. (1945). New Nearctic crane-flies (Tipulidae, Diptera), Part XXVI. Canadian Entomologist 77, 204–208.
| New Nearctic crane-flies (Tipulidae, Diptera), Part XXVI.Crossref | GoogleScholarGoogle Scholar |
Alexander, C. P. (1949). New or little-known Tipulidae (Diptera). LXXXIV. Oriental-Australasian species. Annals & Magazine of Natural History 2, 431–454.
| New or little-known Tipulidae (Diptera). LXXXIV. Oriental-Australasian species.Crossref | GoogleScholarGoogle Scholar |
Alexander, C. P. (1950a). New or little-known Tipulidae (Diptera), LXXXVI. Oriental-Australasian species. Annals & Magazine of Natural History 3, 301–324.
| New or little-known Tipulidae (Diptera), LXXXVI. Oriental-Australasian species.Crossref | GoogleScholarGoogle Scholar |
Alexander, C. P. (1950b). Undescribed species of Japanese crane-flies (Diptera: Tipulidae). Part VII. Annals of the Entomological Society of America 43, 418–436.
| Undescribed species of Japanese crane-flies (Diptera: Tipulidae). Part VII.Crossref | GoogleScholarGoogle Scholar |
Alexander, C. P. (1952). Entomological results from the Swedish expedition 1934 to Burma and British India. Arkiv för Zoologi 3, 123–135.
Alexander, C. P. (1953). Records and descriptions of Japanese Tipulidae (Diptera), Part II. The crane-flies of Shikoku, II. Philippine Journal of Science 82, 21–75.
Alexander, C. P. (1954). Records and descriptions of Japanese Tipulidae (Diptera). Part III. The crane-flies of Shikoku. III. Philippine Journal of Science 82, 263–308.
Alexander, C. P. (1957). Records and descriptions of Japanese Tipulidae (Diptera). Part V. The crane-flies of Honshu. I. Philippine Journal of Science 85, 349–388.
Alexander, C. P. (1958). Records and descriptions of Japanese Tipulidae (Diptera), Part VI. The crane-flies of Honshu, II. Philippine Journal of Science 86, 281–330.
Alexander, C. P. (1959). Undescribed species of crane-flies from the Himalaya Mountains (Tipulidae, Diptera). III. Journal of the New York Entomological Society 66, 161–170.
Alexander, C. P. (1962). New or little-known Tipulidae from eastern Asia (Diptera). XLIX. Philippine Journal of Science 90, 155–214.
Alexander, C. P. (1964). Undescribed species of Nearctic Tipulidae (Diptera), V. The Great Basin Naturalist 24, 19–25.
Alexander, C. P. (1966a). Undescribed species of crane flies from the Himalaya Mountains (Diptera: Tipulidae). XIII. Journal of the New York Entomological Society 74, 180–184.
Alexander, C. P. (1966b). New subgenera and species of crane-flies from California (Ptychopteridae and Tipulidae: Diptera). Transactions of the American Entomological Society 92, 103–132.
Alexander, C. P. (1967). The crane flies of California. Bulletin of the California Insect Survey 8, 1–263.
Alexander, C. P. (1971). Undescribed species of crane flies from the Himalaya Mountains (Diptera: Tipulidae). XIX. Journal of the New York Entomological Society 78, 201–205.
Alexander, C. P. (1975). New or little-known crane flies from Iran. III. (Diptera: Tipulidae). Journal of the New York Entomological Society 83, 121–128.
Alexander, C. P., and Byers, G. W. (1981). Tipulidae. In ‘Manual of Nearctic Diptera. Vol. 1’. (Eds J. F. McAlpine, B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth and D. M. Wood) pp. 153–190. (Research Branch, Agriculture Canada: Ottawa.)
Barnes, H. F. (1937). Methods of investigating the bionomics of the common crane fly, Tipula paludosa Meigen, together with some results. Annals of Applied Biology 24, 356–368.
| Methods of investigating the bionomics of the common crane fly, Tipula paludosa Meigen, together with some results.Crossref | GoogleScholarGoogle Scholar |
Bertone, M. A., Courtney, G. W., and Wiegmann, B. M. (2008). Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes. Systematic Entomology 33, 668–687.
| Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes.Crossref | GoogleScholarGoogle Scholar |
Bremer, K. (1994). Branch support and tree stability. Cladistics 10, 295–304.
| Branch support and tree stability.Crossref | GoogleScholarGoogle Scholar |
Brikiatis, L. (2014). The De Geer, Thulean and Beringia routes: key concepts for understanding early Cenozoic biogeography. Journal of Biogeography 41, 1036–1054.
| The De Geer, Thulean and Beringia routes: key concepts for understanding early Cenozoic biogeography.Crossref | GoogleScholarGoogle Scholar |
Chatterjee, S., Goswami, A., and Scotese, C. R. (2013). The longest voyage: tectonic, magmatic, and paleoclimatic evolution of the Indian plate during its northward flight from Gondwana to Asia. Gondwana Research 23, 238–267.
| The longest voyage: tectonic, magmatic, and paleoclimatic evolution of the Indian plate during its northward flight from Gondwana to Asia.Crossref | GoogleScholarGoogle Scholar |
De Jong, H., and Ciliberti, P. (2014). How cold-adapted flightless flies dispersed over the northern hemisphere: phylogeny and biogeography of the snow fly genus Chionea Dalman (Diptera: Limoniidae). Systematic Entomology 39, 563–589.
| How cold-adapted flightless flies dispersed over the northern hemisphere: phylogeny and biogeography of the snow fly genus Chionea Dalman (Diptera: Limoniidae).Crossref | GoogleScholarGoogle Scholar |
De Jong, H., Oosterbroek, P., Gelhaus, J., Reusch, H., and Young, C. (2008). Global diversity of craneflies (Insecta, Diptera: Tipulidea or Tipulidae sensu lato) in freshwater. Hydrobiologia 595, 457–467.
| Global diversity of craneflies (Insecta, Diptera: Tipulidea or Tipulidae sensu lato) in freshwater.Crossref | GoogleScholarGoogle Scholar |
de Queiroz, K. (1998). The general lineage concept of species, species criteria, and the process of speciation: a conceptual unification and terminological recommendations. In ‘Endless Forms: Species and Speciation’. (Eds D. J. Howard and S. H. Berlocher.) pp. 57–75. (Oxford University Press: Oxford, England.)
de Queiroz, K. (2005). Different species problems and their resolution. BioEssays 27, 1263–1269.
| Different species problems and their resolution.Crossref | GoogleScholarGoogle Scholar | 16299765PubMed |
Dienske, J. W. (1987). An illustrated key to the genera and subgenera of the western Palaerctic Limoniidae (Insecta: Diptera), including a description of the external morphology. Stuttgarter Beitrage zur Naturkunde. Serie A, Biologie 409, 1–52.
Doane, R. W. (1900). New North American Tipulidae. Journal of the New York Entomological Society 8, 190.
Dudley, T. L. (1982). ‘Population and production ecology of Lipsothrix spp. (Diptera: Tipulidae).’ Master’s Thesis, Oregon State University, Corvalis.
Dudley, T. L., and Anderson, N. H. (1987). The biology and life cycles of Lipsothrix spp. (Diptera: Tipulidae) inhabiting wood in Western Oregon streams. Freshwater Biology 17, 437–451.
| The biology and life cycles of Lipsothrix spp. (Diptera: Tipulidae) inhabiting wood in Western Oregon streams.Crossref | GoogleScholarGoogle Scholar |
Edwards, F. W. (1938). British short-palped craneflies. Taxonomy of adults. Transactions of the Society for British Entomology 5, 1–168.
Egger, J. (1863). Dipterologische Beitrage. Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 13, 1101–1110.
Fitch, W. M. (1971). Toward defining the course of evolution: minimum change for a specified tree topology. Systematic Zoology 20, 406–416.
| Toward defining the course of evolution: minimum change for a specified tree topology.Crossref | GoogleScholarGoogle Scholar |
Godfrey, A. (2000). Survey for the cranefly Lipsothrix nigristigma. English Nature Research Report 351, 1–33.
Godfrey, A. (2001). Survey for the cranefly Lipsothrix nigristigma in 2000. English Nature Research Report 410, 1–48.
Godfrey, A. (2003). A review of the invertebrate interest of woody debris in England. English Nature Research Reports 513, 1–50.
Goetghebuer, M., and Tonnoir, A. L. (1921). Catalogue raisonne des Tipulidae de Belgique (Suite). Bulletin de la Societe Entomologique de Belgique 3, 47–58.
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 |
Greenwood, D. R., and Wing, S. L. (1995). Eocene continental climates and latitudinal temperature gradients. Geology 23, 1044–1048.
| Eocene continental climates and latitudinal temperature gradients.Crossref | GoogleScholarGoogle Scholar |
Hancock, E. G., Hewitt, S. M., Godfrey, A., and Mullin, E. (2009). Thoracic spiracular gill structure of Lipsothrix (Diptera, Limoniidae) in Britain described from scanning electron micrographs. Zoosymposia 3, 77–87.
Hinton, H. E. (1955). The structure of the spiracular gill of the genus Lipsothrix (Tipulidae), with some observations on the living epithelium isolated in the gill at the pupa-adult molt. Proceedings of the Royal Entomological Society of London. Series A, General Entomology 30, 1–14.
Hood, G. (2000). ‘PopTools-software for analysis of ecological models, Version 2.1.’ Available at www.poptools.org [Accessed September 2005]
Hynes, C. D. (1965). The immature stages of the genus Lipsothrix in the western United States (Diptera, Tipulidae). The Pan-Pacific Entomologist 41, 165–172.
Kotrba, M. (2000). Morphology and terminology of the female postabdomen. In ‘Contributions to a Manual of Palaearctic Diptera, Vol 1’. (Eds L. Papp and B. Darvas.) pp. 75–84. (Science Herald: Budapest.)
Krivosheina, M. G., and Krivosheina, N. P. (2011). To the biology of the limoniid Lipsothrix nobilis (Diptera, Limoniidae) with description of the immature stages. Zoologicheskij Zhurnal 80, 103–1092.
Krzeminski, W. (2001). First record of the genus Lipsothrix Loew (Diptera; Limoniidae) in Baltic amber (Upper Eocene). Polskie Pismo Entomologiczne 70, 239–331.
Lance, G. N., and Williams, W. T. (1967). Mixed-data classificatory programs I.) Agglomerative systems. Australian Computer Journal 1, 15–20.
Loew, H. (1873). ‘Beschreibungen Europäischer Dipteren. Von Johann Wilhelm Meigen. Dritter Band. Systematische Beschreibung der Bekannten Europäischen Zweiflügeligen Insecten. Zehnter Theil Oder Vierter Supplementband.’ (H.W. Schmidt: Halle, Germany.) viii + 320 pp. [In German]
Lomolino, M. V., Riddle, B. R., Whittaker, R. J., and Brown, J. H. (2010). ‘Biogeography, 4th Edition.’ (Sinauer Associates: Sunderland, MA.)
Mace, G. M. (2004). The role of taxonomy in species conservation. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences 359, 711–719.
| The role of taxonomy in species conservation.Crossref | GoogleScholarGoogle Scholar | 15253356PubMed |
Mani, M. S. (1974). Biogeographical Evolution in India. In ‘Ecology and Biogeography in India’. (Ed. M. Mani.) pp. 698–724. (Dr. W. Junk Publishers: The Hague, Netherlands.)
McAlpine, J. F. (1981). Morphology and terminology – adults. Chapter 2. In ‘Manual of Nearctic Diptera. Volume 1. Monography No. 27’. (Eds J. F. McAlpine, J. R. Peterson and D. M. Wood.) pp. 9–63. (Research Branch, Agriculture Canada: Ottawa.)
Morley, R. J. (2000). ‘Origin and Evolution of Tropical Rain Forests.’ (John Wiley: Chichester, UK.)
Oosterbroek, P. (2014). ‘Catalogue of the craneflies of the world.’ Available from http:// ip30.eti.uva.nl/ccw/ [Accessed 13 June 2014]
Oosterbroek, P., and Theowald, B. (1991). Phylogeny of the Tipuloidea based on characters of larvae and pupae (Diptera, Nematocera) with an index to the literature except Tipulidae. Tijdschrift voor Entomologie 134, 211–267.
Osten Sacken, C. R. (1860). New genera and species of North American Tipulidae with short palpi, with an attempt at a new classification of the tribe. Proceedings. Academy of Natural Sciences of Philadelphia 1859, 197–254.
Osten Sacken, C. R. (1888). Studies on Tipulidae. Part 2. Review of the published genera of the Tipulidae brevipalpi. Part IV. Berliner Entomologische Zeitschrift 31, 171–236.
| Studies on Tipulidae. Part 2. Review of the published genera of the Tipulidae brevipalpi. Part IV.Crossref | GoogleScholarGoogle Scholar |
Petersen, M. J., Bertone, M. A., Wiegmann, B. M., and Courtney, G. W. (2010). Phylogenetic synthesis of morphological and molecular data reveals new insights into the higher-level classification of Tipuloidea (Diptera). Systematic Entomology 35, 526–545.
| Phylogenetic synthesis of morphological and molecular data reveals new insights into the higher-level classification of Tipuloidea (Diptera).Crossref | GoogleScholarGoogle Scholar |
Pielou, E. C. (1991). ‘After the Ice Age: the Return of Life to Glaciated North America.’ (University of Chicago Press: Chicago, IL.)
Podeniene, V. (2003). ‘Morphology and ecology of the last instar larvae of the crane flies (Diptera, Tipulomorpha) of Lithuania.’ Ph.D. Thesis, Vilnius University, Lithuania.
R Development Core Team (2004). ‘R: a language and environment for statistical computing.’ Available at http://www.Rproject.org [Accessed June 2004]
Ribeiro, G. C. (2008). Phylogeny of the Limnophilinae (Limoniidae) and early evolution of the Tipulomorpha. Invertebrate Systematics 22, 627–694.
| Phylogeny of the Limnophilinae (Limoniidae) and early evolution of the Tipulomorpha.Crossref | GoogleScholarGoogle Scholar |
Ribeiro, G. C., and Eterovic, A. (2011). Neat and clear: 700 species of crane flies (Diptera: Tipulomorpha) link southern South America and Australasia. Systematic Entomology 36, 754–767.
| Neat and clear: 700 species of crane flies (Diptera: Tipulomorpha) link southern South America and Australasia.Crossref | GoogleScholarGoogle Scholar |
Rogers, J. S., and Byers, G. (1956). The ecological distribution, life history, and immature stages of Lipsothrix sylvia (Diptera; Tipulidae). Occasional Papers of the Museum of Zoology, University of Michigan, No. 572.
Rotheray, G. E. (2000). Search for the larva of the priority cranefly Lipsothrix ecucullata (Diptera, Tipulidae). Scottish Natural Heritage Commissioned Report F99AC309 (unpublished report).
Sanmartín, I., Enghoff, H., and Ronquist, F. (2001). Patterns of animal dispersal, vicariance, and diversification in the Holarctic. Biological Society of the Linnean Society 73, 345–390.
| Patterns of animal dispersal, vicariance, and diversification in the Holarctic.Crossref | GoogleScholarGoogle Scholar |
Savchenko, E. N. (1982). Limoniidae: Eriopterinae. Fauna Ukraini 14(3), 335 pp. (Akademia Nauk Ukrainskoi SSR, Naukova Dumka: Kiev, Ukraine.)
Savchenko, E. N. (1989). ‘Limoniidae Fauna of the USSR.’ (Akademia Nauk Ukrainian SSR: Kiev, Ukraine.)
Schmidt, J., Opgenoorth, L., Höll, S., and Bastrop, R. (2012). Into the Himalayan exile: the phylogeography of the ground beetle Ethira clade supports the Tibetan origin of forest-dwelling Himalayan species groups. PLoS One 7, e45482.
| Into the Himalayan exile: the phylogeography of the ground beetle Ethira clade supports the Tibetan origin of forest-dwelling Himalayan species groups.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC38XhsVOmu7rP&md5=ca31105512c1e11e324e303edf57eafaCAS | 23049805PubMed |
Sinclair, B. J. (2000). Morphology and terminology of Diptera male terminalia. In ‘Contributions to a Manual of Palaearctic Diptera, Vol. 1’. (Eds L. Papp and B. Darvas.) pp. 53–74. (Science Herald: Budapest, Hungary.)
Starỳ, J. (1992). Phylogeny and classification of Tipulomorpha, with special emphasis on the family Limoniidae. Acta Zoologica Cracoviensia 35, 11–36.
Starỳ, J. (2003). Revision of European species of the genus Rhabdomastix (Diptera: Limoniidae). Part 1: introduction and subgenus Lurdia subgen. n. European Journal of Entomology 100, 587–608.
| Revision of European species of the genus Rhabdomastix (Diptera: Limoniidae). Part 1: introduction and subgenus Lurdia subgen. n.Crossref | GoogleScholarGoogle Scholar |
Starỳ, J. (2004). Revision of European species of the genus Rhabdomastix (Diptera: Limoniidae). Part 2: Subgenus Rhabdomastix s. str. European Journal of Entomology 101, 657–687.
| Revision of European species of the genus Rhabdomastix (Diptera: Limoniidae). Part 2: Subgenus Rhabdomastix s. str.Crossref | GoogleScholarGoogle Scholar |
Starỳ, J. (2007). Nomenclatural changes in West Palaearctic Limoniidae and Pediciidae (Diptera), II. Casopis Slezskeho Musea v Opava (A) 56, 23–36.
Takhtajan, A. (1986). ‘Floristic Regions of the World.’ (University of California Press: Berkeley, CA.) [Translated by T. J. Crovello & A. Cronquist]
Tangelder, I. R. M. (1988). The biogeography of the holarctic Nephrotoma dorsalis species-group (Diptera, Tipulidae). Beaufortia 38, 1–35.
Tiffney, B. H. (1985). Perspectives on the origin of the between eastern Asia and North America. Journal of the Arnold Arboretum 66, 73–94.
Triska, F. J., and Cromak, K. (1980). The role of wood debris in forests and streams. In ‘Forests: Fresh Perspectives from Ecosystem Analysis, Proceedings of the 40th Ann. Biology Colloquium’. (Ed. R. H. Warning.) (Oregon State University Press: Corvallis, OR.)
UKBP (2008). ‘UK Biodiversity Partnership.’ Available at www.ukbap.org.uk/ [Accessed June 2013]
Walker, F. (1848). ‘List of the Specimens of Dipterous Insects in the Collection of the British Museum.’ British Museum: London.
Ward, J. H., (1963). Hierarchical grouping to optimize an objective function. Journal of the American Statistical Association 58, 236–244.
| Hierarchical grouping to optimize an objective function.Crossref | GoogleScholarGoogle Scholar |
Wen, J. (1999). Evolution of eastern Asian and eastern North American disjunct distributions in flowering plants. Annual Review of Ecology and Systematics 30, 421–455.
| Evolution of eastern Asian and eastern North American disjunct distributions in flowering plants.Crossref | GoogleScholarGoogle Scholar |
Willerslev, E., Davison, J., Moora, M., Zobel, M., Coissac, E., Edwards, M. E., Lorenzen, E. D., Vestergård, M., Gussarova, G., Haile, J., Craine, J., Gielly, L., Boessenkool, S., Epp, L. S., Pearman, P. B., Cheddadi, R., Murray, D., Bråthen, K. A., Yoccoz, N., Binney, H., Cruaud, C., Wincker, P., Goslar, T., Alsos, I. G., Bellemain, E., Brysting, E. K., Elven, R., Sønstebø, J. H., Murton, J., Sher, A., Rasmussen, M., Rønn, R., Mourier, T., Cooper, A., Austin, J., Möller, P., Froese, D., Zazula, G., Pompanon, F., Rioux, D., Niderkorn, V., Tikhonov, A., Savvinov, G., Roberts, R. G., MacPhee, R. D. E., Gilbert, M. T. P., Kjær, K. H., Orlando, L., Brochmann, C., and Taberlet, P. (2014). Fifty thousand years of Arctic vegetation and megafaunal diet. Nature 506, 47–51.
| Fifty thousand years of Arctic vegetation and megafaunal diet.Crossref | GoogleScholarGoogle Scholar | 1:CAS:528:DC%2BC2cXhsl2lsL4%3D&md5=df7f6a72cdc4e27fcbf9429ad5d0bd75CAS | 24499916PubMed |
Wolfe, J. A. (1987). Late Cretaceous-Cenozoic history of deciduousness and the terminal Cretaceous event. Paleobiology 13, 215–226.
Yu, Y., Harris, A. J., and He, X.-J. (2010). S-DIVA (statistical dispersal-vicariance analysis): a tool for inferring biogeographic histories. Molecular Phylogenetics and Evolution 56, 848–850.
| S-DIVA (statistical dispersal-vicariance analysis): a tool for inferring biogeographic histories.Crossref | GoogleScholarGoogle Scholar | 20399277PubMed |
Yu, Y., Harris, A. J., and He, X.-J. (2014). RASP (Reconstruct Ancestral State in Phylogenies) 3.0. Available at http://mnh.scu.edu.cn/soft/blog/RASP [Accessed June 2014]