Phylogeny of Cyttaria inferred from nuclear and mitochondrial sequence and morphological data

Abstract

Cyttaria species (Leotiomycetes, Cyttariales) are obligate, biotrophic associates of Nothofagus (Hamamelididae, Nothofagaceae), the southern beech. As such Cyttaria species are restricted to the southern hemisphere, inhabiting southern South America (Argentina and Chile) and southeastern Australasia (southeastern Australia including Tasmania, and New Zealand). The relationship of Cyttaria to other Leotiomycetes and the relationships among species of Cyttaria were investigated with newly generated sequences of partial nucSSU, nucLSU and mitSSU rRNA, as well as TEF1 sequence data and morphological data. Results found Cyttaria to be defined as a strongly supported clade. There is evidence for a close relationship between Cyttaria and these members of the Helotiales: Cordierites, certain Encoelia spp., Ionomidotis and to a lesser extent Chlorociboria. Order Cyttariales is supported by molecular data, as well as by the unique endostromatic apothecia, lack of chitin and highly specific habit of Cyttaria species. Twelve Cyttaria species are hypothesized, including all 11 currently accepted species plus an undescribed species that accommodates specimens known in New Zealand by the misapplied name C. gunnii, as revealed by molecular data. Thus the name C. gunnii sensu stricto is reserved for specimens occurring on N. cunninghamii in Australia, including Tasmania. Morphological data now support the continued recognition of C. septentrionalis as a species separate from C. gunnii. Three major clades are identified within Cyttaria: one in South America hosted by subgenus Nothofagus, another in South America hosted by subgenera Nothofagus and Lophozonia, and a third in South America and Australasia hosted by subgenus Lophozonia, thus producing a non-monophyletic grade of South American species and a monophyletic clade of Australasian species, including monophyletic Australian and New Zealand clades. Cyttaria species do not sort into clades according to their associations with subgenera Lophozonia and Nothofagus.

• Encoelioideae
• Leotiomycetes
• Nothofagus
• southern hemisphere

We are grateful for the aid of I.J. Gamundí and P.R. Johnston who respectively offered invaluable advice and was a host KRP’s field trips to Argentina and New Zealand. We thank two anonymous reviewers for comments. We also thank M. Rajchenberg, A.I. Romero, M. Soto, C. Barroetaveña, A. Ruffini, T.P. Mengoni, C.C. López Lastra, Administracion de Parques Nacionales (collecting permit) (Argentina); T.W. May, A.M. Young, N. Prakash, N. Fechner, P. George, Parks Victoria (collecting permit 10001623) (Australia); E. Gibellini, J.A. Cooper, D.P. Mahoney, A. Bell, Landcare Research (collecting permit) (New Zealand); M.J. Cafaro, C. Currie, C.D. Bell, R.W. Lichtwardt, D.E. Desjardin, B.A. Perry, E. Macapinlac, K. Griffith, M.M. White, L. Williams, G. Giribet, C.C. Davis (USA). We are grateful for the aid of theses herbaria and their staff who loaned material: AD, BAFC, BCRU, BPI, BRI, BRIP, CANB, CUP, DAR, GB, HO, IMI, K, MEL, NY, PDD, S, SFSU, SGO, TRTC. We thank DUKE for transferring many of the aforementioned loans to FH for our use. We also thank the staff of FH and the HUH for their time and expertise, including that of the library of the Gray Herbarium of Harvard University for permission to reproduce the Darwin illustration. We acknowledge financial support from the Department of Organismic and Evolutionary Biology of Harvard University and the Fernald Fund of Harvard University Herbaria, as well as NSF PEET grant DEB-9521944 to D.H. Pfister and M.J. Donoghue.