Abstract
About 150 species of Puccinia on Carex (Cyperaceae) have been reported globally. However, these species can be difficult to identify due to similarities in morphology and difficulties in identifying the host plants. The presence of many cryptic species is also suspected. Based on about 130 specimens on Carex and about 40 related specimens of spermogonial and aecial stages, all collected in northeastern China, a phylogenetic approach was conducted to clarify their identification and life cycles. In the phylogenetic analyses, 17 clades among specimens on Carex were detected. Each clade was interpreted as an independent species, although many clades were morphologically similar to each other. After comparative morphology with species previously reported, seven species including three first records in China (P. circaeae-caricis, P. urticae-inflatae and P. vaginatae) were identified, while ten species (P. caricis-artemisiae, P. caricis-atractylodes, P. caricis-hebeiensis, P. caricis-jilinensis, P. caricis-lactucae, P. caricis-pediformis, P. caricis-rafaensis, P. caricis-ribicola, P. caricis-tenuiformis and P. caricis-violae) are described as new. In uredinial and telial stages, many host plant species could not be identified as they lacked an inflorescence. Thus, a phylogenetic approach is more suitable for species identifications of Puccinia on Carex rather than relying on rust morphology and host plant identification. Life cycle relations for 12 spermogonial and aecial species were also clarified using phylogenetic analyses. Furthermore, it is suspected that Puccinia species on Carex can be differentiated into species based on their spermogonial and aecial stages on the same or related host plants. Additionally, one new replacement name (P. caricis-aphanolepidis) for P. pulchella is proposed.
References
Abbasi, M., Stephen, B., Goodwin, B. & Scholler, M. (2005) Taxonomy, phylogeny, and distribution of Puccinia graminis, the black stem rust: new insights based on rDNA sequence data. Mycoscience 46: 241–247. https://doi.org/10.1007/s10267-005-0244-x
Aime, M.C. (2006) Toward resolving family-level relationships in rust fungi (Uredinales). Mycoscience 47: 112–122. https://doi.org/10.1007/s10267-006-0281-0
Aime, M.C., Bell, C.D. & Wilson, A.W. (2018) Deconstructing the evolutionary complexity between rust fungi (Pucciniales) and their plant hosts. Studies in Mycology 89: 143–152. https://doi.org/10.1016/j.simyco.2018.02.002
Aime, M.C. & McTaggart, A.R. (2021) A higher-lank classification for rust fungi, with notes on genera. Fungal Systematics and Evolution 7: 21–47. https://doi.org/10.3114/fuse.2021.07.02
Aime, M.C., McTaggart, A.R., Mondo, S.J. & Duplessis, S. (2017) Phylogenetics and phylogenomics of rust fungi. Advances in Genetics 100: 267–307. https://doi.org/10.1016/bs.adgen.2017.09.011
Arthur, J.C. (1934) Manual of the rusts in United States and Canada. Purdue Research Foundation, USA.
Azbukina, Z.M. (2005) Rust fungi, Cryptogamic plants, fungi and mosses of the Russian Far East, vol. 5. Dalnauka, Vladivostok, Russia.
Azbukina, Z.M. (2015) Definitorium fungorum Rossiae, Ordo Pucciniales 1. Dal’nauka, Vladivostok, Russia.
Berlin, A., Wallenhammar, A.C. & Andersson, B. (2018) Population differentiation of Puccinia coronata between hosts–implications for the epidemiology of oat crown rust. European Journal of Plant Pathology 152: 901–907. https://doi.org/10.1007/s10658-018-01605-x
Cunningham, G.H. (1931) The rust fungi of New Zealand. John McIndoe Printer, Dunedin, New Zealand.
Cummins, G.B. (1951) Uredinales of continental China collected by S. Y. Cheo II. Mycologia 43: 78–98.
Cummins, G.B. & Hiratsuka, Y. (2003) Illustrated genera of rust fungi, 3<sup>rd</sup> ed. American Phytopathological Society, St. Paul, Minnesota, USA.
Dixon, L.J., Castlebury, L.A., Aime, M.C., Glynn, N.C. & Comstock, J.C. (2010) Phylogenetic relationships of sugarcane rust fungi. Mycological Progress 9: 459–468. https://doi.org/10.1007/s11557-009-0649-6
Engkhaninun, J., Ono, Y. & Kakishima, M. (2005) Phylogenetic relationships of four Puccinia species parasitic on Artemisia in Japan. Mycoscience 46: 61–65. https://doi.org/10.1007/S10267-004-0213-9
Gäumann, E. (1959) Die Rostpilze Mitteleuropas. Buchdruckerei Bucheler Co, Bern, Germany.
Harada, Y. (1977) Puccinia caricis-circaearum, sp. nov., heteroecious Carex rust fungus from Japan. Transactions of Mycological Society of Japan 18: 375–380.
Harada, Y. (1984) Materials for the rust flora of Japan IV. Transactions of Mycological Society of Japan 25: 287–294.
Harada, Y. (1986) Four new species of heteroecious Puccinia (Uredinales) on Carex from Japan. Transactions of Mycological Society of Japan 27: 357–370.
Hasler, A. (1930) Beitrage zur Kenntnis einiger Carex-Puccinien. Annales Mycologici 28: 345–357.
Hiratsuka, N. (1960) A provisional list of Uredinales of Japan proper and the Ryukyu Islands. The Science Bulletin of the Division of Agriculture, Home Economics and Engineering, University of Ryukyus 7: 189–314.
Hiratsuka, N. & Chen, Z.C. (1991) A list of Uredinales collected from Taiwan. Transactions of Mycological Society of Japan 32: 3–22.
Hiratsuka, N., Sato, S., Katsuya, K., Kakishima, M., Hiratsuka, Y., Kaneko, S., Ono, Y., Sato, T., Harada, Y., Hiratsuka, T. & Nakayama, K. (1992) The rust flora of Japan. Tsukuba-shuppankai, Tsukuba, Japan.
Hiratsuka, Y. & Sato, S. (1982) Morphology and taxonomy of rust fungi. In: Scott, K.J. & Chakravorty, A.K. (Eds.) The rust fungi. Academic Press, London, UK, pp. 1–36.
Hrab?tová, M., Kola?ík, M. & Marková, J. (2015) Phylogeny and taxonomy of grass rusts with aecia on Ranunculus and Ficaria. Mycological Progress 14: 12. https://doi.org/10.1007/s11557-015-1033-3
Hylander, N., Jørstad, I. & Nannfeldt, J.A. (1953) Enumeratio Uredinearum Scandinavicarum. Opera Botany 1: 1–102.
Ito, S. (1950) Mycological flora of Japan, vol. 2, no. 3. Yokendo, Tokyo, Japan.
Ito, S. & Homma, Y. (1938) Notae mycologicae Asiae orientalis 3. Tranzactions Sapporo Natural History Society 15: 113–128.
Ji, J.X., Li, Z., Wan, Q., Li, Y. & Kakishima, M. (2016) Notes on rust fungi in China 1. Autoecious life cycle of Puccinia tatarinovii on Prenanthes. Mycotaxon 131: 653–661.
Ji, J.X., Li, Z., Wan, Q., Li, Y. & Kakishima, M. (2017) Notes on rust fungi in China 3. Puccinia adenocauli comb. nov. and its life cycle and new host. Mycotaxon 132: 141–148.
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019a) Two new species of Pucciniastrum producing dimorphic sori and spores from northeast of China. Mycological Progress 18: 529–540. https://doi.org/10.1007/s11557-018-1460-z
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019b) Life cycle of Nothoravenelia japonica and its phylogenetic position in Pucciniales, with special reference to the genus Phakopsora. Mycological Progress 18: 855–864. https://doi.org/10.1007/s11557-019-01496-0
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019c) Notes on rust fungi in China 7. Aecidium caulophylli life cycle inferred from phylogenetic evidence and renamed as Puccinia caulophylli comb. nov. Mycotaxon 134: 719–730. https://doi.org/10.5248/134.719
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020a) Notes on rust fungi in China 8. Pucciniastrum tiliae life cycle and new host plants inferred from phylogenetic evidence. Mycotaxon 135: 491–500. https://doi.org/10.5248/135.491
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020b) Life cycle and taxonomy of Chrysomyxa succinea in China and phylogenetic positions of Caeoma species on Rhododendron. Forest Pathology 50: e12585. https://doi.org/10.1111/efp.12585
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020c) Life cycle and taxonomy of Melampsora abietis-populi in China and its phylogenetic position in Melampsora on Populus. Mycological Progress 19: 1281–1291. https://doi.org/10.1007/s11557-020-01624-1
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2022) Phylogenetic approach for identification and life cycles of Puccinia (Pucciniaceae) species on Poaceae from northeastern China. Phytotaxa 533 (1): 1–48. https://doi.org/10.11646/phytotaxa.533.1.1
Jørstad, I. (1954) The rust on Cyperaceae and Iridaceae in Norway. Skrifter Norske Vidensk Akademie 1954: 1–28.
Juel, O. (1893) Om nagra heteroeciska Uredineer. Botaniska Notiser 1892-1893: 51–57.
Kakishima, M., Yokoi, M. & Harada, Y. (1999) Puccinia carici-adenocauli, a new rust fungus on Carex, and its anamorph, Aecidium adenocauli. Mycoscience 40: 503–507.
Kakishima, M. & Sato, S. (1980) Three Puccinia species on Carex. Transactions of Mycological Society of Japan 21: 35–45.
Kakishima, M. & Sato, S. (1983) Puccinia kawakamiensis, a new caricicolous rust, produces the aecial state on Circaea erubescens. Transactions of Mycological Society of Japan 24: 403–408.
Kenaley, S.C., Hudler, G.W. & Bergstrom, G.C. (2016) Detection and phylogenetic relationships of Puccinia emaculata and Uromyces graminicola (Pucciniales) on switchgrass in New York State using rDNA sequence information. Fungal Biology 120: 791–806. https://doi.org/10.1016/j.funbio.2016.01.016
Kern, F.D. (1917) Noth American species of Puccinia on Carex. Mycologia 9: 205–238.
Kim, C.J. (1963) A provisional list of Uredinales of Korea. Korean Journal of Microbiology 1: 51–64.
Leppik, E.E. (1965) Some viewpoints on the phylogeny of rust fungi. V. Evolution of biological specialization. Mycologia 57: 6–22.
Leppik, E.E. (1967) Some viewpoints on the phylogeny of rust fungi. VI. Biogenic radiation. Mycologia 59: 568–579.
Liu, M. & Hambleton, S. (2010) Taxonomic study of stripe rust, Puccinia striiformis sensu lato, based on molecular and morphological evidence. Fungal Biology 114: 881–899.
Liu, M. & Hambleton, S. (2013) Laying the formation for a taxonomic review of Puccinia coronata s.l. in a phylogenetic context. Mycological Progress 12: 63–89. https://doi.org/10.1007/s11557-012-0814-1
Maier, W., Wingfield, B.D., Mennicken, M. & Wingfield, M.J. (2007) Polyphyly and two emerging lineages in the rust genera Puccinia and Uromyces. Mycological Research 111: 176–185. https://doi.org//10.1016/j.mycres.2006.11.005
McTaggart, A.R., Shivas. R.G., Van der Nest, M.A., Roux, J., Wingfield, B.D. & Wingfield, M.J. (2016) Host jumps shaped the diversity of extant rust fungi (Pucciniales). New Phytologist 209: 1149–1158. https://doi.org/10.1111/nph.13686
Miura, M. (1928) Flora of Manchuria and east Mongolia ?. Cryptogams, Fungi. Minamimanshutetsudo, Dalian, China.
Ono, Y. (1983) Aecial stage of Puccinia dioicae var. micropuncta, var. nov. occurs on Artemisia in Japan. Transactions of Mycological Society of Japan 24: 1–6.
Ono, Y. (1995) Aecidium alangii is the aecial state of Puccinia miyakei (Uredinales). Nova Hedwigia 60: 157–164.
Ono, Y. (2008) Why life-cycle studies?: Implications in the taxonomy of rust fungi (Uredinales). Nippon Kingakukai Kaiho 49: 1–28. https://doi.org/10.18962/jjom.jjom.H19-01
Ono, Y. & Kakishima, M. (1981) Puccinia pulchella: a new Viola-Carex rust from Japan. Canadian Journal of Botany 59: 1543–1546.
Padamsee, M. & McKenzie, E.H.C. (2017) The intriguing and convoluted life of a heteroecious rust fungus in New Zealand. Plant Pathology 66: 1248–1257. https://doi.org/10.1111/ppa.12672
Peck, C.H. (1873) Descriptions of new species of fungi. Bulletin of the Buffalo Society of Natural Sciences 1: 41–72.
Petersen, R.H. (1974) The rust fungus life cycle. Botanical Review 40: 453–513.
Rambaut, A. (2014) FigTree v1.4.2, A graphical viewer of phylogenetic trees. [https://tree.bio.ed.ac.uk/software/figtree/]
Roy, B.A., Vogler, D.R., Bruns, T.D. & Szaro, T.M. (1998) Cryptic species in the Puccinia monoica complex. Mycologia 90: 846–853. https://doi.org/10.1080/00275514.1998.12026978
Sato, T. & Sato, S. (1982) Aeciospore surface structure of the Uredinales. Transactions of Mycological Society of Japan 23: 51–63.
Sato, T. & Sato, S. (1985) Morphology of aecia of the rust fungi. Transactions of British Mycological Society 85: 223–238.
Semmouri, I., Bauters, K., Léveillé-Bourret, É., Starr, J.R., Goetghebeur, P. & Larridon, I. (2019) Phylogeny and systematics of Cyperaceae, the evolution and importance of embryo morphology. Botanical Review 85: 1–39.
Spalink, D., Pender, J., Escudero, M., Hipp, A.L., Roalson, E.H., Starr, J.R., Waterway, M.J., Bohs, L. & Sytsma, K.J. (2018) The spatial structure of phylogenetic and functional diversity in the United States and Canada: An example using the sedge family (Cyperaceae). Journal of Systematics and Evolution 56: 449–465.
Szabo, L. (2006) Deciphering species complexes: Puccinia andropogonis and Puccinia coronata, examples of differing modes of speciation. Mycoscience 47: 130–136.
Tai, F.L. (1979) Sylloge fungorum Sinicorum. Science Press, Beijing, China.
Taylor, J.W., Jacobson, D.J., Kroken, S., Kasuga, T., Geiser, D.M., Hibbett, D.S. & Fischer, M.C. (2000) Phylogenetic species recognition and species concepts in fungi. Fungal Genetics and Biology 31: 21–32.
Termorshuizen, A.J. & Swertz, C.A. (2011) Dutch rust fungi. Aad Termorsguizen, Netherland.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten.https://doi.org/10.12705/Code.2018
Van der Merwe, M.M., Walker, J., Ericson, L. & Burdon, J.J. (2008) Coevolution with higher taxonomic host groups within the Puccinia/Uromyces rust lineage obscured by host jumps. Mycological Research 112: 1387–1408. https://doi.org/10.1016/j.mycres.2008.06.027
Wang, Y.C. (1951) Index Uredinearum Sinensium. Academia Sinica, Beijing, China.
Wilson, M. & Henderson, D.M. (1966) The British rust fungi. Cambridge University Press, Cambridge, UK.
Zambino, P.J. & Szabo, L.J. (1993) Phylogenetic relationships of selected cereal and grass rusts based on rDNA sequence analysis. Mycologia 85: 401–414. https://doi.org/10.1080/00275514.1993.12026292
Zhao, P., Kakishima, M., Wang, Q. & Cai, L. (2017) Resolving the Melampsora epitea complex. Mycologia 109: 391–407. http://dx.doi.org/10.1080/00275514.2017.1326791
Zhao, P., Wang, Q.H., Tian, C.M. & Kakishima, M. (2015) Integrating a numerical taxonomic method and molecular phylogeny for species delimitation of Melampsora species (Melampsoraceae, Pucciniales) on willows in China. PLoS ONE 10: e0144883.https://doi.org/10.1371/journal.pone.0144883
Zhuang, J.Y., Wei, S.X. & Wang, Y.C. (1998) Flora fungorum Sinicorum, vol.10, Uredinales (1). Science Press, Beijing, China.
Aime, M.C. (2006) Toward resolving family-level relationships in rust fungi (Uredinales). Mycoscience 47: 112–122. https://doi.org/10.1007/s10267-006-0281-0
Aime, M.C., Bell, C.D. & Wilson, A.W. (2018) Deconstructing the evolutionary complexity between rust fungi (Pucciniales) and their plant hosts. Studies in Mycology 89: 143–152. https://doi.org/10.1016/j.simyco.2018.02.002
Aime, M.C. & McTaggart, A.R. (2021) A higher-lank classification for rust fungi, with notes on genera. Fungal Systematics and Evolution 7: 21–47. https://doi.org/10.3114/fuse.2021.07.02
Aime, M.C., McTaggart, A.R., Mondo, S.J. & Duplessis, S. (2017) Phylogenetics and phylogenomics of rust fungi. Advances in Genetics 100: 267–307. https://doi.org/10.1016/bs.adgen.2017.09.011
Arthur, J.C. (1934) Manual of the rusts in United States and Canada. Purdue Research Foundation, USA.
Azbukina, Z.M. (2005) Rust fungi, Cryptogamic plants, fungi and mosses of the Russian Far East, vol. 5. Dalnauka, Vladivostok, Russia.
Azbukina, Z.M. (2015) Definitorium fungorum Rossiae, Ordo Pucciniales 1. Dal’nauka, Vladivostok, Russia.
Berlin, A., Wallenhammar, A.C. & Andersson, B. (2018) Population differentiation of Puccinia coronata between hosts–implications for the epidemiology of oat crown rust. European Journal of Plant Pathology 152: 901–907. https://doi.org/10.1007/s10658-018-01605-x
Cunningham, G.H. (1931) The rust fungi of New Zealand. John McIndoe Printer, Dunedin, New Zealand.
Cummins, G.B. (1951) Uredinales of continental China collected by S. Y. Cheo II. Mycologia 43: 78–98.
Cummins, G.B. & Hiratsuka, Y. (2003) Illustrated genera of rust fungi, 3<sup>rd</sup> ed. American Phytopathological Society, St. Paul, Minnesota, USA.
Dixon, L.J., Castlebury, L.A., Aime, M.C., Glynn, N.C. & Comstock, J.C. (2010) Phylogenetic relationships of sugarcane rust fungi. Mycological Progress 9: 459–468. https://doi.org/10.1007/s11557-009-0649-6
Engkhaninun, J., Ono, Y. & Kakishima, M. (2005) Phylogenetic relationships of four Puccinia species parasitic on Artemisia in Japan. Mycoscience 46: 61–65. https://doi.org/10.1007/S10267-004-0213-9
Gäumann, E. (1959) Die Rostpilze Mitteleuropas. Buchdruckerei Bucheler Co, Bern, Germany.
Harada, Y. (1977) Puccinia caricis-circaearum, sp. nov., heteroecious Carex rust fungus from Japan. Transactions of Mycological Society of Japan 18: 375–380.
Harada, Y. (1984) Materials for the rust flora of Japan IV. Transactions of Mycological Society of Japan 25: 287–294.
Harada, Y. (1986) Four new species of heteroecious Puccinia (Uredinales) on Carex from Japan. Transactions of Mycological Society of Japan 27: 357–370.
Hasler, A. (1930) Beitrage zur Kenntnis einiger Carex-Puccinien. Annales Mycologici 28: 345–357.
Hiratsuka, N. (1960) A provisional list of Uredinales of Japan proper and the Ryukyu Islands. The Science Bulletin of the Division of Agriculture, Home Economics and Engineering, University of Ryukyus 7: 189–314.
Hiratsuka, N. & Chen, Z.C. (1991) A list of Uredinales collected from Taiwan. Transactions of Mycological Society of Japan 32: 3–22.
Hiratsuka, N., Sato, S., Katsuya, K., Kakishima, M., Hiratsuka, Y., Kaneko, S., Ono, Y., Sato, T., Harada, Y., Hiratsuka, T. & Nakayama, K. (1992) The rust flora of Japan. Tsukuba-shuppankai, Tsukuba, Japan.
Hiratsuka, Y. & Sato, S. (1982) Morphology and taxonomy of rust fungi. In: Scott, K.J. & Chakravorty, A.K. (Eds.) The rust fungi. Academic Press, London, UK, pp. 1–36.
Hrab?tová, M., Kola?ík, M. & Marková, J. (2015) Phylogeny and taxonomy of grass rusts with aecia on Ranunculus and Ficaria. Mycological Progress 14: 12. https://doi.org/10.1007/s11557-015-1033-3
Hylander, N., Jørstad, I. & Nannfeldt, J.A. (1953) Enumeratio Uredinearum Scandinavicarum. Opera Botany 1: 1–102.
Ito, S. (1950) Mycological flora of Japan, vol. 2, no. 3. Yokendo, Tokyo, Japan.
Ito, S. & Homma, Y. (1938) Notae mycologicae Asiae orientalis 3. Tranzactions Sapporo Natural History Society 15: 113–128.
Ji, J.X., Li, Z., Wan, Q., Li, Y. & Kakishima, M. (2016) Notes on rust fungi in China 1. Autoecious life cycle of Puccinia tatarinovii on Prenanthes. Mycotaxon 131: 653–661.
Ji, J.X., Li, Z., Wan, Q., Li, Y. & Kakishima, M. (2017) Notes on rust fungi in China 3. Puccinia adenocauli comb. nov. and its life cycle and new host. Mycotaxon 132: 141–148.
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019a) Two new species of Pucciniastrum producing dimorphic sori and spores from northeast of China. Mycological Progress 18: 529–540. https://doi.org/10.1007/s11557-018-1460-z
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019b) Life cycle of Nothoravenelia japonica and its phylogenetic position in Pucciniales, with special reference to the genus Phakopsora. Mycological Progress 18: 855–864. https://doi.org/10.1007/s11557-019-01496-0
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2019c) Notes on rust fungi in China 7. Aecidium caulophylli life cycle inferred from phylogenetic evidence and renamed as Puccinia caulophylli comb. nov. Mycotaxon 134: 719–730. https://doi.org/10.5248/134.719
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020a) Notes on rust fungi in China 8. Pucciniastrum tiliae life cycle and new host plants inferred from phylogenetic evidence. Mycotaxon 135: 491–500. https://doi.org/10.5248/135.491
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020b) Life cycle and taxonomy of Chrysomyxa succinea in China and phylogenetic positions of Caeoma species on Rhododendron. Forest Pathology 50: e12585. https://doi.org/10.1111/efp.12585
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2020c) Life cycle and taxonomy of Melampsora abietis-populi in China and its phylogenetic position in Melampsora on Populus. Mycological Progress 19: 1281–1291. https://doi.org/10.1007/s11557-020-01624-1
Ji, J.X., Li, Z., Li, Y. & Kakishima, M. (2022) Phylogenetic approach for identification and life cycles of Puccinia (Pucciniaceae) species on Poaceae from northeastern China. Phytotaxa 533 (1): 1–48. https://doi.org/10.11646/phytotaxa.533.1.1
Jørstad, I. (1954) The rust on Cyperaceae and Iridaceae in Norway. Skrifter Norske Vidensk Akademie 1954: 1–28.
Juel, O. (1893) Om nagra heteroeciska Uredineer. Botaniska Notiser 1892-1893: 51–57.
Kakishima, M., Yokoi, M. & Harada, Y. (1999) Puccinia carici-adenocauli, a new rust fungus on Carex, and its anamorph, Aecidium adenocauli. Mycoscience 40: 503–507.
Kakishima, M. & Sato, S. (1980) Three Puccinia species on Carex. Transactions of Mycological Society of Japan 21: 35–45.
Kakishima, M. & Sato, S. (1983) Puccinia kawakamiensis, a new caricicolous rust, produces the aecial state on Circaea erubescens. Transactions of Mycological Society of Japan 24: 403–408.
Kenaley, S.C., Hudler, G.W. & Bergstrom, G.C. (2016) Detection and phylogenetic relationships of Puccinia emaculata and Uromyces graminicola (Pucciniales) on switchgrass in New York State using rDNA sequence information. Fungal Biology 120: 791–806. https://doi.org/10.1016/j.funbio.2016.01.016
Kern, F.D. (1917) Noth American species of Puccinia on Carex. Mycologia 9: 205–238.
Kim, C.J. (1963) A provisional list of Uredinales of Korea. Korean Journal of Microbiology 1: 51–64.
Leppik, E.E. (1965) Some viewpoints on the phylogeny of rust fungi. V. Evolution of biological specialization. Mycologia 57: 6–22.
Leppik, E.E. (1967) Some viewpoints on the phylogeny of rust fungi. VI. Biogenic radiation. Mycologia 59: 568–579.
Liu, M. & Hambleton, S. (2010) Taxonomic study of stripe rust, Puccinia striiformis sensu lato, based on molecular and morphological evidence. Fungal Biology 114: 881–899.
Liu, M. & Hambleton, S. (2013) Laying the formation for a taxonomic review of Puccinia coronata s.l. in a phylogenetic context. Mycological Progress 12: 63–89. https://doi.org/10.1007/s11557-012-0814-1
Maier, W., Wingfield, B.D., Mennicken, M. & Wingfield, M.J. (2007) Polyphyly and two emerging lineages in the rust genera Puccinia and Uromyces. Mycological Research 111: 176–185. https://doi.org//10.1016/j.mycres.2006.11.005
McTaggart, A.R., Shivas. R.G., Van der Nest, M.A., Roux, J., Wingfield, B.D. & Wingfield, M.J. (2016) Host jumps shaped the diversity of extant rust fungi (Pucciniales). New Phytologist 209: 1149–1158. https://doi.org/10.1111/nph.13686
Miura, M. (1928) Flora of Manchuria and east Mongolia ?. Cryptogams, Fungi. Minamimanshutetsudo, Dalian, China.
Ono, Y. (1983) Aecial stage of Puccinia dioicae var. micropuncta, var. nov. occurs on Artemisia in Japan. Transactions of Mycological Society of Japan 24: 1–6.
Ono, Y. (1995) Aecidium alangii is the aecial state of Puccinia miyakei (Uredinales). Nova Hedwigia 60: 157–164.
Ono, Y. (2008) Why life-cycle studies?: Implications in the taxonomy of rust fungi (Uredinales). Nippon Kingakukai Kaiho 49: 1–28. https://doi.org/10.18962/jjom.jjom.H19-01
Ono, Y. & Kakishima, M. (1981) Puccinia pulchella: a new Viola-Carex rust from Japan. Canadian Journal of Botany 59: 1543–1546.
Padamsee, M. & McKenzie, E.H.C. (2017) The intriguing and convoluted life of a heteroecious rust fungus in New Zealand. Plant Pathology 66: 1248–1257. https://doi.org/10.1111/ppa.12672
Peck, C.H. (1873) Descriptions of new species of fungi. Bulletin of the Buffalo Society of Natural Sciences 1: 41–72.
Petersen, R.H. (1974) The rust fungus life cycle. Botanical Review 40: 453–513.
Rambaut, A. (2014) FigTree v1.4.2, A graphical viewer of phylogenetic trees. [https://tree.bio.ed.ac.uk/software/figtree/]
Roy, B.A., Vogler, D.R., Bruns, T.D. & Szaro, T.M. (1998) Cryptic species in the Puccinia monoica complex. Mycologia 90: 846–853. https://doi.org/10.1080/00275514.1998.12026978
Sato, T. & Sato, S. (1982) Aeciospore surface structure of the Uredinales. Transactions of Mycological Society of Japan 23: 51–63.
Sato, T. & Sato, S. (1985) Morphology of aecia of the rust fungi. Transactions of British Mycological Society 85: 223–238.
Semmouri, I., Bauters, K., Léveillé-Bourret, É., Starr, J.R., Goetghebeur, P. & Larridon, I. (2019) Phylogeny and systematics of Cyperaceae, the evolution and importance of embryo morphology. Botanical Review 85: 1–39.
Spalink, D., Pender, J., Escudero, M., Hipp, A.L., Roalson, E.H., Starr, J.R., Waterway, M.J., Bohs, L. & Sytsma, K.J. (2018) The spatial structure of phylogenetic and functional diversity in the United States and Canada: An example using the sedge family (Cyperaceae). Journal of Systematics and Evolution 56: 449–465.
Szabo, L. (2006) Deciphering species complexes: Puccinia andropogonis and Puccinia coronata, examples of differing modes of speciation. Mycoscience 47: 130–136.
Tai, F.L. (1979) Sylloge fungorum Sinicorum. Science Press, Beijing, China.
Taylor, J.W., Jacobson, D.J., Kroken, S., Kasuga, T., Geiser, D.M., Hibbett, D.S. & Fischer, M.C. (2000) Phylogenetic species recognition and species concepts in fungi. Fungal Genetics and Biology 31: 21–32.
Termorshuizen, A.J. & Swertz, C.A. (2011) Dutch rust fungi. Aad Termorsguizen, Netherland.
Turland, N.J., Wiersema, J.H., Barrie, F.R., Greuter, W., Hawksworth, D.L., Herendeen, P.S., Knapp, S., Kusber, W.-H., Li, D.-Z., Marhold, K., May, T.W., McNeill, J., Monro, A.M., Prado, J., Price, M.J. & Smith, G.F. (2018) International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Koeltz Botanical Books, Glashütten.https://doi.org/10.12705/Code.2018
Van der Merwe, M.M., Walker, J., Ericson, L. & Burdon, J.J. (2008) Coevolution with higher taxonomic host groups within the Puccinia/Uromyces rust lineage obscured by host jumps. Mycological Research 112: 1387–1408. https://doi.org/10.1016/j.mycres.2008.06.027
Wang, Y.C. (1951) Index Uredinearum Sinensium. Academia Sinica, Beijing, China.
Wilson, M. & Henderson, D.M. (1966) The British rust fungi. Cambridge University Press, Cambridge, UK.
Zambino, P.J. & Szabo, L.J. (1993) Phylogenetic relationships of selected cereal and grass rusts based on rDNA sequence analysis. Mycologia 85: 401–414. https://doi.org/10.1080/00275514.1993.12026292
Zhao, P., Kakishima, M., Wang, Q. & Cai, L. (2017) Resolving the Melampsora epitea complex. Mycologia 109: 391–407. http://dx.doi.org/10.1080/00275514.2017.1326791
Zhao, P., Wang, Q.H., Tian, C.M. & Kakishima, M. (2015) Integrating a numerical taxonomic method and molecular phylogeny for species delimitation of Melampsora species (Melampsoraceae, Pucciniales) on willows in China. PLoS ONE 10: e0144883.https://doi.org/10.1371/journal.pone.0144883
Zhuang, J.Y., Wei, S.X. & Wang, Y.C. (1998) Flora fungorum Sinicorum, vol.10, Uredinales (1). Science Press, Beijing, China.