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Rust fungi (Pucciniales) in Changbaishan and
the surrounding area, Jilin, China
Yue-Tai WENG * Yu XUE*, Yoshitaka ONO **
(Accepted November 1, 2016)
Abstract
Changbaishan is the highest mountain complex in northeast China and the Korean Peninsula
located at 41°41' to 42°51'N and 127°43' to 128°16'E on the border between P. R. China and D. P.
R. Korea. Despite well-acknowledged geological and biological diversity in Changbaishan and the
surrounding area, fungal diversity was poorly explored. We studied rust fungi collected in the area
in 2015 and those listed in literature. This paper reports 120 species in 18 genera of the rust fungi
distributed in the area.
Keywords – Geographic distribution
・Northeast Asia
・
Rust fungus diversity
・
Taxonomy
Introduction
Changbaishan (in Chinese; Paektu-san in North Korean and Baekdu-san in South Korean) is
located at 41°41’49’’N – 42°51’18’’N and 127°42’55’’E – 128°16’48’’E (http://www.unesco.org/new/
en/ natural-sciences/environment/ecological-sciences/biosphere-reserves/asia-and-the-pacific/china/
changbaishan/, accessed 5 October 2016) on the border between P. R. China and D. P. R. Korea, and
in Antu County, Jilin Province, P. R. China (Fig. 1). Because it is of volcanic origin with repeated
eruptions in the past and the most recent major eruption in 1903, it is composed of several peaks, each
exceeding 2500 m above sea level (Fig. 2A), surrounding the crater lake, Heaven Lake. The highest peak
is 2744 m above sea level. Changbaishan is the highest mountain complex in northeast China and the
Korean Peninsula. Changbaishan exhibits unique geological formation and is covered with characteristic
vegetation (Fig. 2B–D). It has been designated as one of the MBA Biosphere Reserves by UNESCO
(http://www.unesco.org/new/en/natural-sciences/environment/ sciences/ biosphere-reserves/asia-and-the-
pacic/china/changbaishan/, accessed 5 October 2016).
Climatically, the Changbaishan area is classied in the Xiaoshan Da Hinggan Ling, Changbai Shan
Department of Forestry, School of Forestry, Northeast Forestry University, Harbin, Heilongjian, 150040, P. R. China
Department of Science Education, College of Education, Mito, 310-8215, Japan
*
**
Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）19 －35
20 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
mid-temperate, humid zone (mean January temperature –22.5°C, mean July temperature 21.0°C, annual
precipitation 625 mm: Zheng et al. 2010, Zhang and Zheng 2015). In the phytogeographical classication,
this area is central to the Northeast China Region of the Sino-Japanese Subkingdom, the Eastern Asiatic
Kingdom (Takhtajan 1986, Sun and Wu 2015), where Pinus koraiensis Siebold & Zucc. is a representative
tree, and Astilboides and Mundenia (both Saxifragaceae) and Plagiorhegma (Berberidaceae), Brachybotrys
(Boraginaceae), Diplandrorchis (Orchidaceae) and Symphyllocarpus (Asteraceae) are characteristic
components among others. The flora of this region is composed of species of temperate characteristics,
containing 1776 species of seed plants in 575 genera, 116 families. The ora of this region has a close afnity
to that of northern Japan.
Fig. 1. Location of Changbaishan and the study area in Changbaishan National Nature Reserve and
Ardao Baihe in Antu County, Jilin Province, China. The area surveyed in 2015 is encircled by solid line.
The vegetation of the Changbaishan area falls in the category of the temperate mixed coniferous
and deciduous broad-leaved forest zones (Fig. 2D; Li et al. 2001, Chen 2015). Pinus koraiensis dominates
together with Acer pictum Thunb. subsp. mono (Maxim.) H. Ohashi, Fraxinus mandshurica Rupr.,
Phellodendron amurense Rupr., Populus maximowiczii A. Henry var. barbinervis Nakai (= P. ussuriensis
Kom.), and Tilia amurensis Rupr. var. amureisis in this vegetation zone. Betula platyphylla Skaczev, Quercus
mongolica Fisch., Populus spp., Pinus sylvestris L. var. sylvestris and Larix olgensis A. Henry occur in the
secondary forest where P. koraiensis has been removed. Due to the high elevation of Changbaishan, five
vegetation types are recognized according to the difference in the elevation: Betula costata–Pinus koraiensis–
Abies holophylla–Ulmus forests at 700-1100 m; Pinus koraiensis–Abies nephrolepis forests at 1100-1400 m;
21
Wen et al.: Rust fungi in Changbaishan
Fig. 2. Vegetation of Changbaishan. A. Peaks constituting the summit of Changbaishan above Picea
jezoensis–Abies nephrolepis forests. B. Alpine maedow at ca. 2400 m. C. Picea jezoensis–Abies
nephrolepis forests at ca. 1600 m. D. Mixed coniferous and deciduous broad-leaved forest at ca. 1000 m.
Picea jezoensis–Abies nephrolepis forests at 1400-1800 m (Fig. 2C); Betula ermani eln forest at 1800-2100
m; and small shrub-moss and herbaceous alpine meadow/tundra above 2100 m (Fig. 2B).
Despite well-acknowledged geological and biological diversity in Changbaishan and the surrounding
area, fungal diversity was poorly explored. We studied rust fungi collected in the area in 2015 and those listed
in literature. This paper reports 120 species in 18 genera distributed in the Changbaishan area.
22 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
Materials and Methods
Field survey and specimen collection were undertaken in Changbaishan National Nature Reserve
and the Ardau Baihe area (surrounded by solid line in Fig. 1) in September 8–12, 2015. The rust fungus
specimens collected in the eld survey were deposited in Department of Forestry, Northeast Forestry
University and the Herbarium of Systematic Mycology, College of Education, Ibaraki University (IBAR).
The specimens are listed under each rust fungus taxon in the Result and Discussion section. The fungus
names accepted in Index Fungorum (http://www.indexfungorum.org, accessed 26 September 2016) were
adopted wherever appropriate in this study. Similarly, scientific names listed in Yonekura and Kajita
(2003–. BG plants vernacular name–scientic name index (YList). http:// ylist.info/index.html, accessed
26 September 2016) were used wherever applicable. Names of collectors were abbreviated in initials:
YX (Yue-Tai Wen), YW (Yu Xue) and YO (Yoshitaka Ono).
Microscopic slides were prepared using the methods described by Ono et al. (2012) and Pota et
al. (2013). Sorus structure and spore morphology were observed with an Olympus BX51 microscope
(Olympus, Tokyo, Japan) and bright-eld and differential interference contrast (DIC) optics. Morphological
characteristics observed by microscopy and host identity were compared to those described for individual
species in the flora fungorum sinicorum (Zhuang et al. 1998, 2003, 2005, 2012), the rust flora of Japan
(Hiratsuka et al. 1992) and original descriptions, wherever necessary, for the fungus identication. Published
records of the Changbaishan rust fungi were compiled from Xue (1993) and Zhuang et al. (1998, 2003, 2005,
2012). In the Results and Discussion section, rust genera were arranged according to a taxonomic system
proposed by Cummins and Hiratsuka (2003). Rust species were listed in alphabetical order under each genus.
Map in Fig. 1 were prepared with free blank maps downloaded from “d-maps.com” at http://d-maps.com/
pays.php?num_pay=517&lang=en (accessed 26 September 2016).
Results and Discussion
Pucciniastraceae
Hyalopsora aspidiotus (Peck) Magnus on Gymnocarpium dryopteris (L.) Newman (Cystopteridaceae):
Changbaishan, North Trail, Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11097); on Thelypteris
phegopteris (L.) Sloss. ex Rydb. (Thelypteridaceae): Changbaishan, Xue 1993: 36 (the host listed as
Phegopteris polypodioides).
Hyalopsora hakodatensis Hirats. f. on Cornopteris sp. (Athyriaceae), Changbaishan, Xue 1993: 36.
Hyalopsora polypodii (Dietel) Magnus on Athyrium yokoscense (Franch. & Sav.) Christ (Athyriaceae):
Changbaishan, North Trail, Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11092).
Melampsoridium alni (Thüm.) Dietel on Alnus mandschurica (Callier) Hand.-Mazz. (Betulaceae): Changbaishan,
Xue 1993: 27, 30 & 36; Changbaishan, North Trail, on a trail to waterfall, 8 Sep 2015. YX, YW & YO (IBAR11023);
on S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11057).
23
Wen et al.: Rust fungi in Changbaishan
Melampsoridium betulinum (Pers.) Kleb. on Betula ermanii Cham. (Betulaceae): Changbaishan, Xue 1993:
36 & 38 (listed as Melampsoridium betulae): on Betula platyphylla Sukaczev: Changbaishan, Xue 1993: 27,
30 & 33 (listed as Melampsoridium betulae).
Naohidemyces vaccinii (Jørst.) S. Sato, Katsuya & Y. Hirats. ex Vanderweyen & Fraiture on Vaccinium
uliginosum L. (Ericaceae): Changbaishan, Xue 1993: 38 (listed as Thekopsora vaccinii); on Vaccinium vitis-
idaea L.: Changbaishan, Xue 1993: 38 (listed as Thekopsora vaccinii).
Pucciniastrum agrimoniae (Dietel) Tranzschel on Agrimonia pilosa Ledeb. (Rosaceae): Changbaishan Xue
1993: 28, 31 & 33; East side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11051).
Pucciniastrum areolatum (Fr.) G.H. Otth on Padus racemosa Lam. (Rosaceae): Changbaishan, on S208 to
the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11062); on Picea jezoensis
(Siebold & Zucc.) Carriere (Pinaceae): Changbaishan, Xue 1993: 36 (listed as Thekopsora areolata).
Pucciniastrum asterum (Tranzschel) Jørst. on Aster pekinensis (Hance) F.H. Chen (Asteraceae):
Changbaishan, Xue 1993: 31 (listed as Thekopsora asterum on Kalimeris integrifolia).
Pucciniastrum brachybotrydis (Tranzschel) Jørst. on Brachybotryis paridiformis Maxim. ex Oliv.
(Boraginaceae): Changbaishan, on S208 to the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX,
YW & YO (IBAR11081); Changbaishan, Xue 1993: 28 & 31 (listed as Thekopsora brachybotrydis).
Pucciniastrum epilobii (Pers.) G.H. Otth. on Chamerion angustifolium (L.) Holub. (Onagraceae):
Changbaishan, Xue 1993: 36.; Changbaishan, North Trail, 8 Sep 2015. YX, YW & YO (IBAR11020); North
Trail, on a trail to waterfall, 8 Sep 2015. YX, YW & YO (IBAR11025); North Trail, Underground Forest, 12
Sep 2015. YX, YW & YO (IBAR11089); on S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep
2015. YX, YW & YO (IBAR11063 & 11066).
Pucciniastrum goeppertianum (J.G. Kühn.) Kleb. on Vaccinium vitis-idaea L. (Ericaceae): Changbaishan,
Xue 1993: 35 (listed as Calyptospora goeppertiana).
Pucciniastrum guttatum (J. Schröt.) Hyl., Jørst. & Nannf. on Galium manshuricum Kitag. (Rubiaceae):
Changbaishan, Xue 1993: 30 (listed as Thekopsora guttata).
Pucciniastrum rubiae (Kom.) Jørst. on Rubia argyi (H. Lév. & Vaniot) H. Hara ex Lauener (Rubiaceae): East
side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11050); Changbaishan, on S208 to the South Gate,
ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11056); Changbaishan, Xue 1993: 31 (listed
as Thekopsora rubiae).
Pucciniastrum tiliae Miyabe on Tilia amurensis Rupr. var amurensis (Malvaceae): Ardao Baihe, the
Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11094); Changbaishan, on S208 to the
South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11082); Changbaishan, Xue
1993: 31 & 34.
Uredinopsis licina (Niessl) Magnus on Thelypteris phegopteris (L.) Sloss. ex Rydb. (Thelypteridaceae):
Changbaishan, North Trail, on a trail to Emerald Lake, 8 Sep 2015. YX, YW & YO (IBAR11026); North
Trail, Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11086); Changbaishan, on S208 to the
South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11076); on Thelypteris
sp.: Changbaishan, North Trail, on a trail to waterfall, 8 Sep 2015. YX, YW & YO (IBAR11022).
24 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
Uredinopsis ossiformis Kamei on Dryopteris expansa (C. Presl.) Fraser-Jenk. & Jermy
(Dryopteridaceae): Changbaishan, Xue 1993: 37.
Uredinopsis pteridis Dietel & Holw. on Pteridium aquilinum (L.) Kuhn (Dennstaedtiaceae):
Changbaishan, Xue 1993: 28 & 33.
Uredinopsis struthiopteridis F.C.M. Störmer on Matteuccia struthiopteris (L.) Tod. (Onocleaceae):
Changbaishan, Xue 1993: 37.
Fig. 2. Uredinium and spores of Changbaishan rust fungi. A. Amphispores of Hyalopsora aspidiotus
(IBAR11097). B. Urediniospores of Pucciniastrum areolatum (IBAR11062). C. Peridiate uredinium
of Pucciniastrum brachybotrydis (IBAR11081). D. Urediniospores of Pucciniastrum brachybotrydis
(IBAR11081). E. Uredinium of Uredinopsis filicina (IBAR11022). F. Amphispores of Uredinopsis
licina (IBAR11022). Bars: 20 µm in A, B, D–F; 100 µm in C.
25
Wen et al.: Rust fungi in Changbaishan
Fig. 3. Sori and spores of Changbaishan rust fungi.. A. Aeciospores of Chrysomyxa succinea (IBAR11030).
B. One-celled teliospores and four-celled basidia of Coleosporium tussilaginis (IBAR11043). C. Uredinial
paraphyses of Aplopsora lonicerae (IBAR11078). D. Teliospores of Aplopsora lonicerae (IBAR11078).
E. Basidium and basidiospores of Aplopsora lonicerae (IBAR11078). F. Uredinium of Melampsora
epitea (IBAR11041). Note thick-walled, capitate paraphyses and echinulate uredinispores. G. Telium of
Melampsora epitea (IBAR11041). Note evenly thin-walled teliospores. Bars: 20 µm in A–G.
Coleosporiaceae
Chrysomyxa succinea (Sacc.) Tranzschel on Picea jezoensis Carr. (Pinaceae): Changbaishan, North Trail, on
a trail to Underground Forest, 8 Sep 2015. YX, YW & YO (IBAR11030); on Rhododendron aureum Georgi
(Ericaceae), Changbaishan, Xue 1993: 35 & 38 (listed as Chrysomyxa ledi var. rhododendri); Changbaishan,
South Trail to Heaven Lake, alt. 2100-2200 m, 9 Sep 2015. YX, YW & YO (IBAR11035).
Coleosporium cimicifugatum Thüm. on Actaea spicata H. Hara (Ranunculaceae): Changbaishan, Xue
26 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
1993: 26, 29 & 35; on Cimicifuga simplex (DC.) Wormsk. ex Turcz. (Ranunculaceae): Changbaishan,
Xue 1993: 26 & 29; Changbaishan, on S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep
2015. YX, YW & YO (IBAR11055).
Coleosporium clematidis Barclay on Clematis terniflora DC. var. mandshurica (Rupr.) Ohwi
(Ranunculaceae): Changbaishan, Xue 1993: 26 & 32 (the host listed as Clematis mandshurica).
Coleosporium neocacaliae Saho on Parasenecio hastatus (L.) H. Koyama (Asteraceae): Changbaishan,
on S208 to the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11083);
on Parasenecio auriculatus (DC.) J. R. Grant: Changbaishan, Xue 1993: 35 (listed as Coleosporium
cacaliae on Cacalia auriculata); on Parasenecio kamtschaticus (Maxim.) Kadota: Changbaishan, Xue
1993: 38 (listed as Coleosporium cacaliae on Cacalia kamtschatica).
Coleosporium phellodendri Kom. on Phellodendron amurense Rupr. (Rutaceae), Changbaishan, Xue 1993:
26 & 29; Ardao Baihe, the Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11093); Ardao
Baihe, on S208 to the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11072).
Coleosporium pini-asteris Orish. on Aster scaber Thunb. (Asteraceae): Ardao Baihe, on S208 to the
South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR1080); Ardao Baihe, on
S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11069).
Coleosporium saussureae Thüm. on Saussurea grandifolia Maxim. (Asteraceae), Changbaishan, Xue
1993: 27 & 29.; on Saussurea pulchella (Fisch. ex. Hornem.) Fisch., Changbaishan, Xue 1993: 27, 29,
32 & 33; East side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11053); On S208 to the South
Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11054).
Coleosporium tussilaginis (Per.) Lév. on Adenophora pereskiifolia (Fisch. ex Roem. & Schult.) Fisch.
ex G. Don (Campanulaceae): Changbaishan, Xue 1993: 32 (listed as Coleosporium campanulae); on
Campanula glomerata L. subsp. glomerata (Campanulaceae): Changbaishan, Xue 1993: 32 (listed as
Coleosporium campanulae); on Sonchus oleraceus L. (Asteraceae): East side of Ardao Baihe, 10 Sep
2015. YX, YW & YO (IBAR11043); Ardao Baihe, the Northeast Botanical Garden, 12 Sep 2015. YX,
YW & YO (IBAR11103); on Senecio cannabifolius Less. var. daurica (Asteraceae): Changbaishan, Xue
1993: 27 & 29 (listed as Coleosporium senecionis).
Coleosporium yamabense Saho on Petasites rubellus (J.F. Gmel) Toman (Asteraceae): Changbaishan,
Xue 1993: 38 (listed as Coleosporium petasitis on Petasites saxatilis).
Cronartiaceae
Cronartium accidum (Alb. & Schwein.) G. Winter on Paeonia japonica (Makino) Miyabe & Takeda
(Paeoniaceae): Ardao Baihe, the Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11104).
Cronartium ribicola J.C. Fisch. on Ribes mandshuricum (Maxim.) Kom. (Grossulariaceae):
Changbaishan, Xue 1993: 27.
Melampsoraceae
Melampsora caprearum Thüm. on Salix capraea L. (Salicaceae), Changbaishan, North Trail,
27
Wen et al.: Rust fungi in Changbaishan
Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11091); on Salix integra Thunb.:
Changbaishan, Xue 1993: 30 (listed as Melampsora laricis-caprearum); on Salix sp. (Salix cf. futura
Seemen): Changbaishan, North Trail, on a trail to waterfall, 8 Sep 2015. YX, YW & YO (IBAR11024).
Melampsora epiphylla Dietel on Salix nummularia Andersson (Salicaceae): Changbaishan, Xue 1993:
38 (the host listed as Salix rotundifolia); on Salix rorida Lalsch.: Changbaishan, on S208 to the South
Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR1060 & 11068).
Melampsora epitaea Thüm. on Salix cf. divaricata Pall. var. metaformosa (Nakai) Kitagawa (Salicaceae):
Changbaishan, North Trail, on a trail to Emerald Lake, 8 Sep 2015. YX, YW & YO (IBAR 11027);
South Trail to Heaven Lake, alt. 2100-2200m, 9 Sep 2015. YX, YW & YO (IBAR1037, 11038 & 11039);
on Salix integra Thunb.: Changbaishan, Xue 1993: 30; on Salix nummularia Andersson: Changbaishan,
South Trail to Heaven Lake, alt. ca. 2400m, 9 Sep 2015. YX, YW & YO (IBAR11041); on Salix
viminalis L.: Ardao Baihe, the Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11098);
on Salix sp., Changbaishan, North Trail, on a trail to Underground Forest, 8 Sep 2015. YX, YW & YO
(IBAR11034); East side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11044); Changbaishan, on
S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11061).
Melampsora itoana (Hirats. f.) S. Ito & Homma on Oxalis acetosella L. (Oxalidaceae): Changbaishan,
on S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR1077); on
Oxalis grifthii Edgew. & Hook. f.: Changbaishan, Xue 1993: 35.
Melampsora laricis-populina Kleb. on Populus maximowiczii A. Henry var. barbinervis Nakai (Salicaceae),
Changbaishan, Xue 1993: 29, 33 (the host as Populus ussuriensis); on Populus nigra L., East side of Ardao
Baihe, 10 Sep 2015. YX, YW & YO (IBAR1046); Populus simonii Carriere, Changbaishan, Xue 1993: 29; on
Populus suaveolens Fischer, Changbaishan, North Trail, on a trail to waterfall, 8 Sep 2015. YX, YW & YO
(IBAR11021); East side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11049); Changbaishan, on S208
to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11059).
Melampsora magnusiana G.H. Wagner on Populus alba L. (Salicaceae): East side of Ardao Baihe, 10
Sep 2015. YX, YW & YO (IBAR11042).
Melampsora populnea (Pers.) P. Karst. on Populus tremula L. var. davidiana (Dode) C.K. Schneid.
(Salicaceae): Changbaishan, Xue 1993: 29 & 33 (listed as Melampsora laricis on Populus davidiana).
Melampsora rostrupii G.H. Wagner on Populus cf. candicans Aiton (Salicaceae): Changbaishan, North
Trail, Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11087).
Melampsora salicis-albae Kleb. on Salix rorida Laksch. (Salicaceae): Changbaishan, Xue 1993: 33.
Phakopsoraceae
Pucciniostele mandshurica Dietel on Astilbe chinensis (Maxim.) Maxim. ex Franch. & Sav.
(Saxifragaceae): Changbaishan, Xue 1993: 28 & 31.
Chaconiaceae
Aplopsora lonicerae Tranzschel on Lonicera maximowizcii (Rupr.) Regel (Caprifoliaceae): Changbaishan, on
28 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11078).
Uropyxidaceae
Triphragmiopsis jeffersoniae Naumov on Jeffersonia dubia (Maxim.) Benth. & Hook. f. (Berberidaceae):
Antu, Zhuang et al. 2012: 110.
Triphragmiopsis laricinum (W.F. Chou) F.L. Tai on Larix gmelini (Rupr.) Rupr. (Pinaceae): Antu,
Zhuang et al. 2012: 111; on Larix olgensis A. Henry: East side of Ardao Baihe, 10 Sep 2015. YX, YW &
YO (IBAR11045); Changbaishan, on S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015.
YX, YW & YO (IBAR11065); Changbaishan, Xue 1993: 37; Changbaishan, Zhuang et al. 2012: 111;
Antu, Zhuang et al. 2012: 111.
Phragmidiaceae
Phragmidium fusiforme J. Schröt. on Rosa acicularis Lindl. (Rosaceae): Antu, Zhuang et al. 2012: 146.
Phragmidium griseum Dietel on Rubus arcticus L. (Rosaceae): Changbaishan, Xue 1993: 31 & 34); on
Rubus crataegifolius Bunge: Antu, Zhuang et al. 2012: 174; East side of Ardao Baihe, 10 Sep 2015. YX,
YW & YO (IBAR11047).
Phragmidium kamtschatkae (H.W. Anderson) Arthur & Cummins on Rosa amblyotis C.A. Mey.
(Rosaceae): Changbaishan, Xue 1993: 32, 33 (the host listed as Rosa davurica).
Phragmidium miyakeanum Hirats. on Rubus mesogaeus Focke. (Rosaceae): Changbaishan, on S208 to
the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11075).
Phragmidium montivagum Arthur on Rosa acicularis Lindl. (Rosaceae): Antu, Zhuang et al. 2012: 151.
Phragmidium papillatum Dietel on Potentilla cryptotaeniae Maxim. (Rosaceae): Antu, Zhuang et al.
2012: 138; on Potentilla chinensis Ser.: Changbaishan, Xue 1993: 32.
Phragmidium rosae-multiorae Dietel on Rosa acicularis Lindl. (Rosaceae): Changbaishan, Xue 1993:
36; on Rosa amblyotis C.A. Mey.: Changbaishan, Xue 1993: 28 & 32 (the host listed as Rosa daurica).
Xenodochus carbonarius Schltdl. on Sanguisorba stipulata Raf. (Rosaceae): Changbaishan, South Trail to
Heaven Lake, alt. 2100-2200m, 9 Sep 2015. YX, YW & YO (IBAR11040); on Sanguisorba sitchensis C.A.
Mey.: Antu, Z huang et al. 2012: 196; on Sanguisorba tenuifolia Fisch. ex Link: Antu, Zhuang et al. 2012: 196.
Pucciniaceae
Gymnosporangium clavariiforme Dietel on Crataegus maximowiczii C.K. Schneid. (Rosaceae): Antu,
Zhuang et al. 2012: 8.
Gymnosporangium tremelloides R. Hartig on Sorbus pohuashanensis (Hance) Hedl. (Rosaceae):
Changbaishan, Xue 1993: 35.
Puccinia albulense (Magnus) Kuntze on Veronicastrum sibiricum (L.) Pennell var. sibiricum
(Plantaginaceae): Changbaishan, Xue 1993: 31 (listed as Puccinia veronicarum on Veronica sibirica).
Puccinia alisovae Tranzschel on Angelica sp. (Apiaceae): Antu, Zhuang et al. 2003: 111.
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Wen et al.: Rust fungi in Changbaishan
Puccinia allii (DC.) F. Rudolphi on Allium sp. (Amaryllidaceae): Antu, Zhuang et al. 1998: 243.
Puccinia angelicae (Schumach.) Fuckel on Anagelica anomala Ave-Lall. (Apiaceae): Changbaishan,
Xue 1993: 27, 30 & 32.
Puccinia angelicicola Henn. on Angelica dahurica (Fisch. ex Hoofm.) Benth. & Hook. f. ex Franch. & Sav.
(Apiaceae): Antu, Zhuang et al. 2003: 113.
Puccinia aomoriensis Syd. & P. Syd. on Atractylodes ovata (Thunb.) DC. (Asteraceae): Antu, Zhuang et al.
1998: 179 (the host listed as Atractylodes japonica); Xue 1993: 31 (the host listed as Atractylodes japonica).
Puccinia argentata (Schultz) G. Winter on Impatiens noli-tangere L. (Balsaminaceae): Changbaishan,
Fig. 4. Teliospores of Changbaishan rust fungi. A. Triphragmiopsis laricinum (IBAR11045). B.
Phragmidium miyakeanum (IBAR11075). C. Xenodochus carbonarius (IBAR11040). D. Puccinia
argentata (IBAR11032). E. Puccinia hultenii (IBAR11036). F. Uromyces heimerlianus (IBAR11105).
Bars: 20 µm in A, D–F; 100 µm in B & C.
30 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
North Trail, on a trail to Underground Forest, 8 Sep 2015. YX, YW & YO (IBAR11032).
Puccinia artemisiae-keiskeanae Miura on Artemisia stolonifera (Maxim.) Kom. (Asteraceae): Antu,
Zhuang et al. 2003: 205.
Puccinia arundinellae-anomalae Dietel on Arundinella hirta (Thunb.) Tanaka (Poaceae): Changbaishan,
Xue 1993: 27.
Puccinia asiatica (Kom.) P. Syd. & Syd. on Mitella nuda L. (Saxifragaceae): Changbaishan, Xue 1993:
36; Changbaishan, North Trail, Underground Forest, 12 Sep 2015. YX, YW & YO (IBAR11090).
Puccinia atrofusca (Dudley & C.H. Thomps.) Holw. on Artemisia selengensis Turcz. ex Besser
(Asteraceae): Antu, Zhuang et al. 1998: 179.
Puccinia baicalensis Tranzschel on Aegopodium alpestre Ledeb. (Apiaceae): Antu, Zhuang et al. 2003: 114.
Puccinia behenis G.H. Otth on Gypsophila oldhamiana Miq. (Caryophyllaceae): Antu, Zhuang et al.
2003: 49.
Puccinia brachypodii Otth on Agrostis alba L. (Poaceae): Antu, Zhuang et al. 1998: 53; on
Anthoxanthum odoratum L. subsp. nipponicum (Honda) Tzvelev (Poaceae): Changbaishan, Zhuang et al.
1998: 53 (the host listed as Anthoxanthum nipponicum).
Puccinia bupleuri (Opiz) F. Rudolphi on Bupleurum longiradiatum Turcz. (Apiaceae): Changbaishan,
Xue 1993: 27 & 30.
Puccinia calcitrapae DC. on Cirsium schantarense Trautv. & C.A. Mey. (Asteraceae): Changbaishan,
Xue 1993: 27, 30, 32, 36 & 38 (listed as Puccinia cirsii); on Cirsium sp.: Antu: Zhuang et al. 2003: 210;
Changbaishan, North Trail, on a trail to Emerald Lake, 8 Sep 2015. YX, YW & YO (IBAR11028); North
Trail, on a trail to Underground Forest, 8 Sep 2015. YX, YW & YO (IBAR11033); Changbaishan, on
S208 to the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11079).
Puccinia caricis-rhizopodae Miura on Carex sp. (Cyperaceae): Antu, Zhuang et al. 1998: 198.
Puccinia circaeae Pers. on Circaea alpina L. (Onagraceae): Changbaishan, North Trail, on a trail to
Underground Forest, 8 Sep 2015. YX, YW & YO (IBAR11029); North Trail, Underground Forest, 12
Sep 2015. YX, YW & YO (IBAR11084); North Trail, Underground Forest, 12 Sep 2015. YX, YW & YO
(IBAR11085); on Circaea caulescens Nakai: Antu, Zhuang et al. 2003: 105; Changbaishan, Xue 1993:
36 (the host listed as Circaea alpina var. caulescens).
Puccinia clintoniae-udensis Bubak on Clintonia udensis Trautv. & Mey. (Liliaceae): Changbaishan,
Zhuang et al. 1998: 246.
Puccinia cnici-oleracei Pers. on Artemisia stolonifera (Maxim.) Kom. (Asteraceae): Antu, Zhuang et al. 2003: 212.
Puccinia coronata Corda on Poa pratensis L. (Poaceae): Changbaishan, Xue 1993: 27; on Rhamnus
davurica Pall. (Rhamnaceae): Changbaishan, Xue 1993: 30 (the host listed as Rhamnus ussuriensis);
Antu, Zhuang et al. 1998: 62 (the host listed as Rhamnus ussuriensis); on Rhamnus schneideri Lév. &
Vant. var. manshurica Nakai: Antu, Zhuang et al. 1998: 62; on Roegneria turczaninovii (Drobov) Nevski
(Poaceae): Antu, Zhuang et al. 1998: 64; Roegneria sp.: Antu, Zhuang et al. 1998: 64.
Puccinia dioicae Magnus on Carex sp. (Cyperaceae): Antu, Zhuang et al. 1998: 204.
Puccinia dioscoreae Kom. on Dioscorea nipponica Makino (Dioscoreaceae): Changbaishan, Xue 1993: 30.
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Wen et al.: Rust fungi in Changbaishan
Puccinia elymi Westend. on Elymus sibiricus L. (Poaceae): Antu, Zhuang et al. 1998: 74.
Puccinia glechomatis DC. on Meehania urticifolia (Miq.) Makino (Laminaceae): Changbaishan, on
S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11074).
Puccinia graminis Pers. subsp. graminicola Z. Urb. on Triticum aestivum L. (Poaceae): Changbaishan,
Xue 1993: 30 (the host listed as Triticum sativum).
Puccinia haleniae Arthur & Holw. on Halenia corniculata (L.) Cornaz. (Gentianaceae): Antu, Zhuang et
al. 2003: 149.
Puccinia helianthi Schwein. on Helianthus annuus L. (Asteraceae): Antu, Zhuang et al. 2003: 221.
Puccinia hieracii (Röhl.) H. Mart. on Taraxacum sp. (Asteraceae): Antu, Zhuang et al. 2003: 224.
Puccinia hultenii Tranzschel & Jørst. on Rumex acetosa L. (Polygonaceae): Changbaishan, South Trail
to Heaven Lake, alt. 2000m, 9 Sep 2015. YX, YW & YO (IBAR11036).
Puccinia lactucae-debilis Dietel on Ixeris chinensis Nakai (Asteraceae): Antu, Zhuang et al. 2003: 228.
Puccinia magnusiana Körn. on Phragmites communis Trin. (Poaceae): Antu, Zhuang et al. 1998: 105.
Puccinia menthae Pers. on Mentha arvensis L. (Laminaceae): Antu, Zhuang et al. 2003: 168; on Mentha
davurica Fisch., Changbaishan, Xue 1993: 31.
Puccinia minussensis Thüm. on Lactuca indica L. (Asteraceae): Antu, Zhuang et al. 2003: 232;
Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11100); on Lactuca triangulata
Maxim.: Changbaishan, Xue 1993: 30; Lactuca sp.: Antu, Zhuang et al. 2003: 233.
Puccinia miyoshiana Dietel on Spodiopogon sibiricus Trin. (Poaceae): Changbaishan, Xue 1993: 30.
Puccinia nanbuana Henn. on Angelica cf. anomala Ave-Lall. (Apiaceae); Changbaishan, on S208 to the
South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11070).
Puccinia okatamaensis S. Ito on Phragmites communis Trin. (Poaceae): Antu, Zhuang et al. 1998: 120.
Puccinia orbicula Peck & Clinton on Nabalus tatarinowii (Maxim.) Nakai (Asteraceae): Changbaishan,
on S208 to the South Gate, ca. 60 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11071).
Puccinia polygoni-amphibii Pers. on Persicaria thunbergii (Siebold & Zucc.) H. Gross (Polygonaceae):
Changbaishan, Xue 1993: 31 (the host listed as Polygonum thunbergii).
Puccinia polygoni-sieboldii (Hirats. f. & S. Kaneko) B. Li on Persicaria sagittata (L.) H. Gross
(Polygonaceae): Antu, Zhuang et al. 2003: 36 (the host listed as Polygonum sieboldii).
Puccinia punctata Link on Galium manshuricum Kitag. (Rubiaceae): Changbaishan, Xue 1993: 27 &
31; Antu, Zhuang et al. 2003: 189.
Puccinia pygmaea Erikss. On Deyeuxia langustifolia (Kom.) Y.L. Chang ex S.L. Lu (Poaceae): Antu,
Zhuang et al. 1998: 142.
Puccinia recondita Dietel & Holw. on Aconitum ochranthum C.A. Mey. (Ranunculaceae): Antu, Zhuang
et al. 1998: 146; on Clematis patens Morr. & Decne. (Ranunculaceae): Antu, Zhuang et al. 1998:
146; on Deyeuxia langsdori (Link) Kunth (Poaceae): Antu, Zhuang et al. 1998: 148; on Thalictrum
aquilegiifolium L. var. sibiricum Regel. & Tiling (Ranunculaceae): Antu, Zhuang et al. 1998: 147.
Puccinia ribis DC. on Ribes komarovii Pojark (Grossulariaceae): Antu, Zhuang et al. 2003: 82; on Ribes
mandshuricum (Maxim.) Kom.: Changbaishan, Xue 1993: 27.
32 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
Puccinia saniculae Grev. on Sanicula rubriora Fr. Schmidt (Apiaceae): Antu, Zhuang et al. 2003: 136.
Puccinia stipae-sibiricae S. Ito on Phedimus aizoon (L.) 't Hart (Crassulaceae): Antu, Zhuang et al.
1998: 161 (the host listed as Sedum aizoon).
Puccinia tanaceti DC. on Artemisia mandshurica Rupr. (Asteraceae): Changbaishan, Xue 1993: 31; on
Artemisia stolonifera (Maxim.) Kom.: Changbaishan, Xue 1993: 34; on Artemisia lavandulaefolia DC.:
Antu, Zhuang et al. 2003: 243.
Puccinia tatarinovii Kom. & Tranzschel on Nabalus tatarinowii (Maxim.) Nakai (Asteraceae):
Changbaishan, Xue 1993: 31 (the host listed as Prenathes tatarinowii).
Puccinia tossoensis Tokun. & Kawai on Synurus deltoides (Aiton) Nakai (Asteraceae): Antu, Zhuang et
al. 2003: 249.
Puccinia violae (Schumach.) DC. on Viola albida Palib. (Violaceae): Antu, Zhuang et al. 2003: 102; on
Viola sacchalinensis Boiss., Changbaishan, North Trail, on a trail to Underground Forest, 8 Sep 2015.
YX, YW & YO (IBAR11031); Ardao Baihe, the Northeast Botanical Garden, 12 Sep 2015. YX, YW
& YO (IBAR11099); Northeast Botanical Garden, 12 Sep 2015. YX, YW & YO (IBAR11101); Viola
verecunda A. Gray: Changbaishan, Xue 1993: 28 & 33.
Puccinia yaramesuge Homma on Carex campylorhiza (Cyperaceae), Changbaishan, Xue 1993: 30.
Uromyces aconiti Fuckel on Aconitum macrorhynchum Turcz. & Ledeb. (Ranunculaceae):
Changbaishan, Xue 1993: 32.
Uromyces amurensis Kom. on Maackia amurensis Rupr. & Maxim. (Fabaceae): Antu, Zhuang et al.
2005: 27; Changbaishan, Xue 1993: 28, 32.
Uromyces appendiculatus F. Strauss on Phaseolus vulgaris L. (Fabaceae): Antu, Zhuang et al. 2005: 30.
Uromyces dactylidis G.H. Otth on Alopecurus aequalis Sobol. (Poaceae): Changbaishan, Zhuang et al.
2005: 117.
Uromyces ervi (Wallr.) Westend. on Vicia pseudo-orobus Fischer & C.A. Mey. (Fabaceae): Changbaishan, on
S208 to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11058); on Vicia
unijuga A. Br.: East side of Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11052).
Uromyces geranii (DC.) G.H. Otth & Wartm. on Geranium krameri Franch. & Sav. (Geraniaceae):
Changbaishan, Xue 1993: 34 (the host listed as Geranium japonicum).
Uromyces hedysari-obscuri (DC.) Carestia & Picc. on Hedysarum vicioides Turcz. subsp. japonicum (B.
Fedtsch.) B.H. Choi & H. Ohashi var. japonicum (B. Fedtsch.) B.H. Choi & Ohashi (Fabaceae): Antu,
Zhuang et al. 2005: 42 (the host listed as Hedysarum vicioides).
Uromyces heimerlianus Magnus on Vicia pseudo-orobus Fischer & C.A. Mey.: Changbaishan, on S208
to the South Gate, ca. 73 km from Ardao Baihe, 11 Sep2015. YX, YW & YO (IBAR11105); on Vicia
unijuga A. Braun: Antu, Zhuang et al. 2005: 43.
Uromyces itoanus Hirats. f. on Kummerowia striata (Thunb.) Schnidl. (Fabaceae): Antu, Zhuang et al.
2005: 53 (listed as Uromyces lespedezae-procumbentis).
Uromyces lespedezae-procumbentis (Schwein.) Lagerh. on Lespedeza bicolor Turcz. (Fabaceae): Antu,
Zhuang et al. 2005: 53; Changbaishan, Xue 1993: 34.
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Wen et al.: Rust fungi in Changbaishan
Uromyces orobi (Schmach.) Lév. on Vicia unijuga A. Braun (Fabaceae): Changbaishan, Xue 1993: 33.
Uromyces sommerfeltii Hyl., Jørst. & Nannf. on Solidago virgaurea L. (Asteraceae): Antu, Zhuang et al.
2005: 106; Changbaishan, Zhuang et al. 2005: 106.
Uromyces striatus J. Schröt. on Trifolium repens L. (Fabaceae): Changbaishan, Xue 1993: 34.
Uromyces trifolii-repentis Liro on Trifolium repens L. (Fabaceae): Changbaishan, on S208 to the South
Gate, ca. 73 km from Ardao Baihe, 11 Sep 2015. YX, YW & YO (IBAR11067).
Uromyces veratri (DC.) J. Schröt. on Veratrum nigrum L. (Melanthiaceae): Antu, Zhuang et al. 2005:
142; Changbaishan, Xue 1993: 32.
Uromyces viciae-fabae (Pers.) J. Schröt. on Lathyrus komarovii Ohwi (Fabaceae): Antu, Zhuang et al.
2005: 73; on Vicia amoena Fisch. ex DC.: Antu, Zhuang et al. 2005: 74; Changbaishan, East side of
Ardao Baihe, 10 Sep 2015. YX, YW & YO (IBAR11048); on Vicia amurensis Oett.: Antu, Zhuang et al.
2005: 74; on Vicia japonica A. Gray: Antu, Zhuang et al. 2005: 74; Vicia cracca L.: Changbaishan, Xue
1993: 28 (listed as Uromyces viciae-craccae); on Vicia unijuga A. Braun: Antu, Zhuang et al. 2005: 74.
Anamorphic Genus
Aecidium viburni Henn. & Shirai on Virburnum sargenti Koehne (Adoxaceae): Changbaishan, Xue
1993: 29.
As to the regional mycodiversity, China is one of geographic regions where rust fungi have been
relatively well studied. Compilation of rust species from the records published by Tai (1979), Teng (1996)
and Zhuang et al. (1998, 2003, 2005, 2012) revealed that 999 species in 57 teleomorphic genera and
76 anamorphic species are distributed in China (Ono unpublished). However, in terms of rust biology,
species distributed in China have been poorly studied. Complete life cycle of 477 species (ca. 47.7 %)
has not been determined. Even in the most species for which the complete life cycle is known, the life
cycle study was carried out with populations of those species outside of China. Seven hundred and sixty-
three species are distributed in Japan; and 351 species (46.0 %) among them remain to be studied for
their complete life cycle (Ono unpublished). However, many species for which the complete life cycle
is known were studied with Japanese populations (Hiratsuka et al. 1984). By contrast, in British Isls,
only 54 species (21.3 %) of a total of 253 species are unknown for their complete life cycle. Taxonomic
identication of the species and elucidation of their biological nature are interactive; and, therefore, a
view of rust diversity (in terms of species richness) will change as the biological study of the species
is advanced. As seen in the above enumeration of species, majority (ca. 80 %) of species recorded in
the Changbaishan area are known from one or two collections. Apparently, more intensive eld survey
and collection are needed for better understanding of the rust fungus diversity in this geologically and
oristically interesting area. As mentioned previously, the Changbaishan area is central to the Northeast
China Region of the Sino-Japanese Subkingdom, the Eastern Asiatic Kingdom (Takhtajan 1986, Sun
and Wu 2015), where 1776 species of seed plants in 575 genera, 116 families are recorded. According
to Rossman (1994), we know only 14 % of a total of 50000 rust species that exist on this globe. By
34 Bull. Coll. Educ., Ibaraki Univ.（Nat. Sci.）66（2017）
analyzing the number of rust species in Neotropics (some Central American countries and Brazil) and
North America (Indiana, USA), Hennen and McCain (1993) assumed that the rate of rust fungi to host
plants in a given area ranges between 1:20 and 1:4. The estimate of 1.5 million fungus species on the
globe was based on an assumed fungus to plant ratio of 6:1 (Hawksworth 1991). If these assumptions are
applicable in the Northeast China Region, 300 or more rust species are expected to be distributed there;
and this gure would be reected to the rust species diversity in the Changbaishan area.
Disclosure
The authors declare no conict of interests. This study complies with the current laws of P. R. China and
Japan.
Acknowledgments
This study was supported, in part, by Grant-in-Aid from Northeast Forestry University. The eld
survey and collecting study materials were carried out under the permission from the Changbaishan
National Nature Protective and Administrative Bureau, Forest Department of Jilin Province, Forest
Ministry of P. R. China.
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