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ALBERT R. MANN LIBRARY
CORNELL UNIVERSITY
CORNELL UNIVERSITY LIBRARY
ili
1 579
YCOTAZXON
AN INTERNATIONAL JOURNAL FOR RESEARCH ON TAXONOMY & NOMENCLATURE OF FUNGI, INCLUDING LICHENS
Volume 90(1) + July-September 2004
CONTENTS
A contribution to the rust flora (Uredinales) of southern Africa, with an emphasis on South Africa Mechthilde Mennicken & Franz Oberwinkler =| New species of Asterina from Guangdong, China Bin Song, Tai-Hui Li & Ya-Heng Shen = 29 Preliminary survey of the Helvellaceae from Xinjuiang, China Wen-ying Zhuang 35 — Genetic variability in tropical and temperate populations of . Trapeliopsis glaucolepidea: Evidence against long range dispersal in a lichen with disjunct distribution 7 Zdenek Palice & Christian Printzen 43 New and rare rust fungi (Uredinales) from Anatolia (Turkey) — 2 Zeliha Bahcecioglu & Halvor B. Gjaerum 55 A new species of Pseudobaeospora from California Dennis E. Desjardin 69 Les pycnides et conidies de Lecanora vaenskaei (lichens, Lecanoraceae) Claude Roux 77 New and previously unrecorded saxicolous species of Buellia s.1. with one-septate ascospores from the Greater Sonoran Desert Region Frank Bungartz 81 Cumminsiella mirabilissima on Mahonia aquifolium in Turkey Elsad Hiiseyin 125 Interesting taxa of Meliolaceae in HMAS, China Bin Song & Tai-Hui Li 129 Molecular analyses confirm the relationship between Stephanospora caroticolor and Lindtneria trachyspora M.P. Martin, S. Raid] & M.T. Telleria 133 A new Neotyphodium species symbiotic with drunken horse grass (Achnatherum inebrians) in China Chunjie Li, Zhibiao Nan, Volker H. Paul, Peter D. Dapprich & Yong Liu 141 Kuehneola uredinis (Uredinales) on species of Rubus in Turkey 5 Elsad Hiiseyin 149 Fungi of alpine ‘islands’ of Dryas octopetala in the Carpathians Andrzej Chlebicki & Markéta Sukova 153
[Content continues inside front cover]
<item ates ae SS oe Se oh cee cere eG Sohn Seabee ee ONE EEE Ue OSE E eer ISSN 0093-4666 MYXNAE 90(1): 1-240 (2004) For subscription details, availability on microform, and availability of articles as photocopies or tear sheets, see back cover
[Content continued from front cover]
Four new lichen species from Turkey Kenan Yazici, Ali Aslan & André Aptroot 177 Nectriella guadalupensis and its Dendrodochium-like anamorph (Bionectriaceae, Hypocreales): anew species on Agavaceae Annette W. Ramaley 181 A host-specific species of Hypoxylon from France, and notes on the chemotaxonomy of the “Hypoxylon rubiginosum complex” Marc Stadler, Hartmund Wollweber & Jacques Fournier 187 Tuber zhongdianense sp. nov. from China Xing Yuan He, Hai Mei Li & Yun Wang 213
Book reviews and notices David L. Hawksworth 217 Penultimate revisions Lorelei L. Norvell 237 Errata . 239
MYCOTAXON
Volume 90, pp.1-2& July-September 2004
A contribution to the rust flora (Uredinales) of southern Africa, with an emphasis on South Africa*
MECHTHILDE MENNICKEN & FRANZ OBERWINKLER
info@ systbot.uni-tuebingen.de Lehrstuhl fiir Spezielle Botanik & Mykologie Botanisches Institut, Universitat Tiibingen Auf der Morgenstelle 1, D-72076 Tiibingen, Germany
Abstract—Six new rust species are described: Puccinia rocherpaniana on Helichrysum tricos- tatum, P. pteroniae on Pteronia divaricata, Uredo tarchonanthi on Tarchonathus littoralis, Uredo aspalathi on Aspalathus laricifolia, Uromyces silksvleyensis on cf. Bartholina burmanniana, and Uromyces quaggafonteinus on Ehrharta calycina. Rust species newly recorded from the western provinces of South Africa and/or on new host plants are: Aecidium elytropappi on Elytropappus rhinocerotis and Leysera gnaphalodes, Aecidium dielsii on Lebeckia sp., Miyagia pseudosphaeria on Sonchus cf. oleraceus, cf. Phakopsora pachyrhizae on Psoralea pinnata, Puccinia polycampta on Chlorophytum crassinerve and on Trachyandra sp., P. kalchbrenneri var. kalchbrenneri on He- lichrysum cochleariforme, P. ursiniae on Ursinia anthemoides, P. granularis on Pelargonium grossularioides, P. menthae on Mentha cf. longifolia, P. hordei on Bromus pectinatus, P. polypog- onis on Polypogon monspeliensis, P. stonemaniae on Thesium cf. strictum and on T. viridifolium, Uredo sp. on Salsola cf. zeyheri, Uromyces chaetobromi on Chaetobromus dregeanus, Uromyces ehrhartae-giganteae (= Uredo ehrhartae-calycinae) on Ehrharta calycina and on E. villosa, Uro- myces holci on cf. Holcus setiger, on Karroochloa cf. schismoides, on Schismus barbatus and on Tribolium echinatum.
Keywords—Biodiversity, Namibia, rust fungi, taxonomy.
Introduction
Up to 400 rust species (Crous, Phillips & Baxter 2000; Doidge 1950) and about 20.000 species of higher plants (Acocks 1953) are known from South Africa. If it is true that rust species are about 5-25% as numerous as plant species (Hennen & McCain 1993), a total rust flora of about 1.000 to 5.000 species could be calculated. Thus, it is likely that a lot more rust species are yet to be found in South Africa.
In the pilot phase of the BIOTA (Biodiversity Monitoring Transect Analyses) Southern Africa project extensive collections of rust fungi were made in the western parts of South Africa and in Namibia. For our first results, we describe six new rust species and we report the first recording of several rust species for South Africa and for Namibia. Furthermore, known rust fungi are shown to have extended host ranges, than currently accepted.
* Part 216 in the series ,,Studies in Heterobasidiomycetes“ from the Botanical Institute, University of Tiibingen.
Materials and methods The examined specimens were mostly collected in the South African BIOTA-observa- tories by the first author and Markus Goker between 17.XI.2001—03.XII.2001 and by the first author between 02.1X.2002-30.IX.2002. The 13 BIOTA-observatories in South Africa were set up along a transect, parallel to the N7, from Cape of Good Hope to Richtersveld. Each observatory has one km’. The specimens collected in the course of this study as well as some additional specimens from herbaria were studied as we per- formed freehand sections and scrape mounts of infected plant material. The samples were heated in “Hoyer’s fluid” (Cunningham 1972) and subsequently examined with a Carl Zeiss microscope with bright field and phase contrast. With the exception of pycnio- and basidiospores, 25 spores of each occurring spore state were measured. If only one collection of a rust species was available 50 spores of each present spore state were measured. The same was done with type collections. If a spore state is put in parentheses, measurements were not available. The reasons for that are either too small a number of spores or strong alterations of spores that had already ger- minated. The cells of the peridium were also examined. In our line drawings, which show the cells as optical sections, the outer wall of the cells is on the left hand side, the inner wall on the right hand side. All specifications with the prefix ‘circa’ are based on measure- ments of less than 25 cells. For the determination of the rust fungi the encyclopaedic publications of Doidge (1927, 1928, 1939, 1941, 1948a, 1948b, 1950) and Cummins (1971) were mostly consulted. In addition, we checked a lot of further publications concerning the rust flora of Africa which are available via internet under http://www.mycology.uni-tuebingen.de/databas- es/rust-literature/.
Results The rust species are listed in alphabetical order referring to the host family and to the host genus, respectively. The following abbreviations are used: 0 = spermogonia = py- cnia; I = aecia; II = uredinia; III = telia; IV = basidia.
ANTHERICACEAE — CHLOROPHYTUM & ASPHODELACEAE —TRACHYANDRA Puccinia polycampta Syd. (Sydow 1924: 236). (Fig. 1)
Type on Chlorophytum sp. (as Anthericum sp.) South Africa, Western Cape Province, Stellen- bosch, [X.1923, leg. L. Verwoerd No. 1260, (Langbaan No. 1270).
Pycnia unknown. Aecia not seen.
Uredinia amphigenous on leaves, sparse in the middle of concentric groups of telia, supepidermal, roundish or ellipsoid in outline, up to 2 mm diameter, dark brown to black, long covered by the epidermis which ruptures lately but always partly veiled, pul- verulent, surrounded by mahogany-red leaf spots. Urediniospores subgloboid, ovoid or ellipsoid, 23-38 x 19-30 um, finely echinulate, spores borne singly on pedicels, spore wall uniformly about 1.5—2 um thick, yellowish to hyaline, germ pores inconspicuous, 8 to 13, scattered, with hyaline papillae that are weakly developed.
Telia amphigenous on leaves, separate or scattered in longitudinal deformed concentric groups, up to 1 cm diameter, partly surrounding one uredinium, subepidermal, dark
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reddish-black, long covered by the epidermis which ruptures lately but always partly veiled, single sori compact, surrounded by mahogany-red leaf spots, loculated by a well developed stroma of dark mahogany-brown, palisade-wise paraphyses. Teliospores in general bicellular, variable in shape and size, ellipsoid, clavate, oblong or irregular in outline, rounded, acuminate or flattened at the apex, rounded or attenuate at the base, slightly constricted at the septum, 38-65 x 18-30 um, spore wall smooth, occasionally with longitudinal surface ridges, about 1—2 um thick at the sides, about 2—8 tm thick at the apex, brown to (pale) chestnut-brown, germ pores obscure, pedicel persistent, up to 20 um long, often truncate short below of the attachment, thin-walled, collapsing, red- dish-brown to hyaline. Mesospores and tricellular spores present.
PEASE REY
Fig. 1: Puccinia polycampta on Chlorophytum crassinerve. Uredinio- and teliospores (RSA 146). Scale bar = 10 um.
Specimens examined:
- On Chlorophytum crassinerve (Bak.) Oberm. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 31.2’’, E 17° 32’ 45.1’’, 11.1X.2002, leg. M. Mennicken No. RSA 146, II Il (PREM, M).
- On Trachyandra sp. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 32.2’, E 18° 18’ 23.3’’, 19.1X.2002, leg. M. Mennicken No. RSA 187, II Ill (PREM, M).
- On Trachyandra sp. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 32.9’, E 18° 18’ 24.0’’, 19.1X.2002, leg. M. Mennicken No. RSA 195, II III (PREM, M).
Specimen not examined because of the scantiness of the infection:
- On Trachyandra sp. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 32.0’, E 18° 18’ 40.1’, 21.1X.2001, leg. M. Mennicken No. RSA 18, III (PREM).
Our collections agree well with the diagnosis of Puccinia polycampta even if the spec- trum of spore measurements is wider in our collections. Sydow (1924) stated for the urediniospores 24—27 x 20-25 um and for the teliospores 36-44 x 18-24 um. Chloro- phytum crassinerve is anew host plant and Puccinia polycampta seems to be new to the rust flora of the Northern Cape Province.
ASTERACEAE - HELICHRYSUM Puccinia kalchbrenneri De-Toni 1888: 645 var. kalchbrenneri. (Fig. 2)
Lectotype on Helichrysum nudifolium Less. South Africa, near Somerset East, Promont, leg. P. MacOwan.
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Syn. Puccinia helichrysi Kalchbrenner & Cooke 1880: 21. Type on Helichrysum petiolatum D. Don, South Africa, No. 35.
Anamorph. Uredo lepisclinis Thiimen 1877: 410. Type on Helichrysum nudifolium. South Africa, near Somerset-East, Promont, 1876, leg. P. MacOwan No. 1239.
Pycnia, aecia, and telia not seen.
Uredinia mostly abaxial on leaves, and on stems, sparse, separate or scattered in irreg- ular groups, roundish to ellipsoid, small, up to 0.5 mm wide, yellowish-brown, exposed, pulverulent. Urediniospores globoid, subgloboid, or ovoid, 22—28 (—31) x 20-25 (—27) um, finely echinulate, spores borne singly on pedicels, spore wall uniformly about 1-2 um thick, bright yellow to hyaline, germ pores inconspicuous, 5 to 9, scattered, without papillae.
Fig. 2: Puccinia kalchbrenneri var. kalchbrenneri on Helichrysum cochleariforme. Uredin- iospores (RSA 196). Scale bar = 10 um.
Specimen examined:
- On Helichrysum cochleariforme DC. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 25.6’’, E 18° 18’ 32.8’’, 19.1X.2002, leg. M. Mennicken No. RSA 196, II (PREM, M).
Our collection conforms well with the description of the uredinia and urediniospores of Puccinia kalchbrenneri given by Gjerum (1998): “Uredinia hypophyllous, yellowish brown. Urediniospores (21—) 25-33 x 17-25 (—29) um, globoid, subgloboid or obovoid, sometimes irregular, wall | um thick, finely echinulate”. Helichrysum cochleariforme seems to be a new host plant.
Puccinia rocherpaniana Mennicken & Oberw. sp. nov. (Fig. 3)
Pycnia et aecia ignota. Uredinia adaxialia, subepidermalia, vulpina, usque ad 1 mm diam., mox nuda, pulveru- lenta. Urediniosporae subgloboideae, ovoideae vel ellipsoideae, interdum angulatae, 25—35 (—38) x 21-30 um, pariete echinulato, aureo-fusco vel olivaceo, 2-3 um crasso, 3-6 poris germinationis dispersis. Telia adaxialia, subepidermalia, nigra, usque ad 1 mm diam., mox nuda, compacta vel pulvinata. Teliosporae bicellulares, obo- voideae, ellipsoideae, clavatae vel oblongae, apice rotundato, acuto vel applanato, base rotundata vel attenuata, medio leviter constrictae, 36-69 um x 21-37 um, pariete laevigato, brunneo vel castaneo, lateraliter 1-2.5 um crasso, apicaliter 4-14 um crasso, poro germinationis cellulae superioris apicali, cellulae inferioris juxta septum posito, pedicello dilute aureo-fusco vel hyalino, usque ad 65 um longo. Mesosporae numerosae, ellipsoideae vel oblongae, apice acuto vel rotundato, base attenuata vel rotundata, 30-46 x 17-26 um, pariete laevi, brunneo vel castaneo, lateraliter 1-2 um crasso, apicaliter 3-11 um crasso, poro germinationis apicali.
In foliis Helichrysi tricostati (Thunb.) Less. (Asteraceae).
Pycnia and aecia unknown.
Uredinia adaxial on leaves, subepidermal, sparse, separate or scattered, roundish in outline, small, up to 1 mm wide, foxy red, exposed, pulverulent, surrounded by the torn epidermis. Urediniospores subgloboid, ovoid or ellipsoid, sometimes irregularly angu- lar, 25-35 (-38) x 21-30 um, finely echinulate, spores borne singly on pedicels, spore
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wall uniformly about 2—3 um thick, brownish-yellow to olive-green, germ pores incon- spicuous 3 to 6, scattered, with hyaline papillae that are weakly developed.
Telia adaxial on leaves, subepidermal, developing from the uredinia, sparse, separate or scattered, black, exposed, compact to cushion-like, surrounded by the torn epidermis. Teliospores in general bicellular, variable in shape and size, obovoid, ellipsoid, clavate or oblong, rounded, acuminate or flattened at the apex, rounded or attenuate at the base, slightly constricted at the septum, 36—69 um x 21—37 um, spore wall smooth, about 1— 2.5 um thick at the sides, about 4-14 um thick at the apex, brown to chestnut-brown, paler towards the base and the apex, germ pores apical and just below the septum, pedicel persistent, up to 65 um long, thin-walled, mostly collapsing, brownish- yellow to hyaline. Mesospores common, ellipsoid to oblong, acuminate or roundish at the apex, attenuate or roundish at the base, 30-46 x 17—26 um, spore wall smooth, about 1—2 um thick at the sides, about 3—11 um thick at the apex, brown to chestnut-brown, paler to- wards the base and the apex, germ pore apical, pedicel similar to the bicellular teliospores.
On the leaves of Helichrysum tricostatum (Asteraceae).
Etymology. Named after the habitat, where the first collection was made, Rocherpan Nature Reserve.
Syisilleis.ee
Fig. 3: Puccinia rocherpaniana on Helichrysum tricostatum. Uredinio- and teliospores (RSA 136). Scale bar = 10 um.
Specimens examined:
- On Helichrysum tricostatum. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 27.0”’, E 17° 32’ 40.1’’, 11.1X.2002, leg. M. Mennicken No. RSA 136, ITI] (Holotype PREM, Isotype M).
- On Helichrysum tricostatum. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 26.9”’, E 17° 32’ 40.1’’, 26.X1.2001, leg. M. Mennicken No. RSA 52, II III (Paratypes PREM, M). - On Helichrysum tricostatum. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 33.3’, E 18° 18’ 36.3’’, 21.X1.2001, leg. M. Mennicken No. RSA 21, IT III (Paratypes PREM, M).
- On Helichrysum tricostatum. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 24.4’, E 18° 18’ 47.9’’, 21.X1.2001, leg. M. Mennicken No. RSA 22, II III (Paratypes PREM, M).
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- On Helichrysum tricostatum. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 32.3’, E 18° 18° 24.7’, 19.1X.2002, leg. M. Mennicken No. RSA 198, II III (Paratypes PREM, M).
As far as could be established, there are five known Puccinia and one Uredo species on the host genus Helichrysum:
The autoecious, macrocyclic Puccinia kalchbrenneri (see above) differs from Puccinia rocherpaniana in having smaller, thinner-walled urediniospores: “20-30 x 19-26 um” (Pole Evans 1915), “(21—) 25-33 x 17-25 (-29) um, ... wall 1 um thick” (Gjerum 1998), and narrower, thinner-walled teliospores having a thinner apex and short, often fragile pedicels: “40-73 x 15-23 un, rarely up to 27 u broad, the majority being 40—57 x 15-22 yt, epispore smooth, delicate, thin, pale except of the apex where it is thickened (up to 10 w) and chestnut brown; ... pedicel short, deciduous” (Doidge 1927), “42-66 x 21-27 um, ... walls pale brown, | um thick, apically darker brown, up to 7 um thick. Pedicels up to 30 um long, often deciduous” (Gjzrum 1998).
Puccinia kalchbrenneri var. valida Doidge 1927: 58 differs from Puccinia rocherpani- ana in having smaller, thinner-walled urediniospores (see above) and in having narrow- er teliospores (50-80 x 18-24 wu) with a deep chestnut brown colouration at the apex. The urediniospores of Puccinia kalchbrenneri var. valida are similar to those of Puc- cinia kalchbrenneri.
Puccinia macowani G. Winter (Winter 1885: 255) and Puccinia pienaarii Pole-Evans (Pole Evans 1915: 643) are autoecious, probably demicyclic rust fungi. Puccinia ma- cowani differs from Puccinia rocherpaniana in having longer and smaller teliospores “60-90 x 19-25 uy” (Doidge 1927). The epispore of the teliospores of Puccinia pienaarii is up to 10 um thick, and has “irregular longitudinal or reticulate ridges or flanges (these is only visible when the spore is dry)” (Doidge 1927).
Puccinia lagenophorae Cooke 1884: 6, which infects several Helichrysum species in Australia (Shivas 1989), is autoecious and demicyclic.
Uredo humbertii Maire 1934: 646, described from Madagascar, has bigger uredin- iospores (40-42 x 34-36 um).
The attributes of Puccinia rocherpaniana are not suitable to any other Puccinia species on Asteraceae cited in Doidge (1927). Therefore, the rust fungus is described as new. The host genus Helichrysum belongs to the tribe Gnaphalieae (Herman et al. 2000). With + 600 species, Helichrysum is widespread in Africa (244 species in southern Af- rica) and Madagascar, as well as in Europe, Asia and Australia.
ASTERACEAE - LEYSERA
Aecidium elytropappi Henn. (Hennings 1898: 294). (Fig. 4)
Type on Elytropappus rhinocerotis Less. South Africa, “West-Capland’, in the hills behind Bot Rivier, XI.1896, leg. R. Schlechter (B 70 0007271).
Pycnia unknown.
Aecia aecidioid, seriate on stems, covering the whole stem surface over several centi- meters, often causing minor hypertrophies of the stems, without leaf spots surrounding aecia, aecial cups conical to cylindrical, erumpent, circa 200-450 um diameter, up to 1 mm long, spore mass freshly orange to yellow, bleaching in the herbarium to cream- white, surrounded by a recurved, irregularly and deeply frayed peridium. Peridium
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freshly orange to yellow, bleaching in the herbarium to cream-white, cells of the perid- ium medium connected, outer wall finely striate, circa 14-16 um wide, inner wall ver- rucous, circa 4—7 um wide. Aeciospores angular globoid, subgloboid to ovoid, 19-27 x 14-22 um, spore wall about 1—2.5 um thick, up to 4 um thick in the edge with the only germ pore, without papillae, finely verrucous, hyaline.
Fig. 4: Aecidium elytropappi on Leysera gnaphalodes. Peridium cells and aeciospores (RSA 200). Scale bar = 10 um.
Specimens examined:
- On Elytropappus rhinocerotis. South Africa, ‘West-Capland’, in the hills behind Bot Rivier, XI.1896, leg. R. Schlechter, I (Type B 70 0007271).
- On Leysera gnaphalodes (L.) L. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Na- ture Reserve, S 32° 36’ 33.3’’, E 18° 18’ 37.0°’, 21.XI.2001, leg. M. Mennicken No. RSA 25, I (PREM, M).
- On Leysera gnaphalodes. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 27.0’, E 18° 18’ 32.2’’, 19.[X.2002, leg. M. Mennicken No. RSA 200, I (PREM, M).
Aecidium elytropappi is denoted on Elytropappus rhinocerotis from the Eastern and Western Cape Province in South Africa (Crous, Phillips & Baxter 2000) and on Stoebe kilimandsharica O. Hoffm. from Kilimanjaro in Tanzania (Henderson 1972). As far as could be established, Leysera is a new host genus. All three known host genera belong to the Gnaphalieae (Herman et al. 2000).
ASTERACEAE - PTERONIA
Puccinia pteroniae Mennicken & Oberw. sp. nov. (Fig. 5)
Pycnia et aecia ignota. Uredinia amphigena, subepidermalia, vulpina, usque ad | mm diam., mox nuda, pulver- ulenta. Urediniosporae subgloboideae, pyriformes, ellipsoideae vel oblongae, saepe angulatae, (28—) 33-41 (— 48) x 21-33 (—36) um, pariete echinulato, aureo-fusco, lateraliter (1.5—) 2—2.5 (—3) um crasso, apicaliter 2-4 um crasso, 24 poris germinationis aequatorialibus. Telia amphigena, subepidermalia, nigra, usque ad 1 mm diam., mox nuda, pulverulenta. Teliosporae bicellulares, subgloboideae, ellipsoideae vel cylindraceae, apice rotundato, base rotundata, medio leviter constrictae, 41-58 (—65) x (28—) 30-40 (-43) um, pariete laevi, lateraliter 3-6 uum crasso, apicaliter usque ad 9 um crasso, bilaminato, strato exteriore tenui, luteo-porphyreo, strato interiore atro-porphyreo, poro germinationis cellulae superioris + apicali vel subapicali, cellulae inferioris + juxta septum posito, pedicello inflato, dilute aureo-fusco vel hyalino, usque ad 95 um longo.
In foliis Pteroniae divaricatae (Berg.) Less. (Asteraceae).
Pycnia and aecia unknown.
Uredinia amphigenous on leaves, subepidermal, sparse, separate or arranged in irregu- lar groups, single sori round, elliptic or irregular in outline, small, up to 1 mm wide, foxy red, early exposed, pulverulent, surrounded by the torn epidermis, with slightly yellowed and/or brightened leaf spots surrounding uredinia. Urediniospores variable in shape and size, subgloboid, pyriform, ellipsoid or oblong, often somewhat angular, (28—) 33-41 (-48) x 21-33 (—36) um, echinulate, echinulation coarsely meshed, spores
8
borne singly on pedicels, spore wall about (1.5—) 2—2.5 (—3) um thick at the sides, about 2—4 um thick at the apex, golden-brown, germ pores conspicuous, 2 to 4, mostly 3, equatorial, with hyaline papillae that are weakly developed.
Telia amphigenous on leaves, subepidermal, developing from the uredinia, sparse, sep- arate or in irregular groups, single sori roundish or irregular in outline, small, up to 1 mm wide, mostly smaller, black, early exposed, pulverulent, surrounded by the torn epidermis, with slightly yellowed and/or brightened leaf spots surrounding telia. Teliospores bicellular, subgloboid, ellipsoid or cylindrical, rounded above and below, slightly constricted at the septum, 41-58 (—65) x (28—) 30-40 (-43) um, spore wall smooth, about 3—6 um thick at the sides, up to 9 um thick at the apex or over the germ pore, bilaminate, outer layer usually thin-walled, forming preferentially in the range of germ pores large, papilla-like swellings, pale and yellowed mahogany-coloured, inner layer + uniformly thick-walled, mahogany-coloured to deep mahogany-coloured, germ pore in the upper cell + apical or 1/3 of the distance from the apex to the septum, in the lower cell below the septum or 1/3 of the distance from the septum to the base, pedicel persistent, up to 95 um long, not collapsing, short below the attachment with cylindri- cal, stripe-like inflation, outer margins of the swelling diffusing in Hoyer’s fluid, brownish-yellow to hyaline, sometimes obliquely inserted.
On the leaves of Pteronia divaricata (Asteraceae).
Etymology. Named after the host plant, Pteronia divaricata.
Fig. 5: Puccinia pteroniae on Pteronia divaricata. Uredinio- and teliospores (RSA 174). Scale bar = 100i.
Specimens examined:
- On Pteronia divaricata. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27’ 43.0’, E 18° 26’ 26.1’’, 18.1X.2002, leg. M. Mennicken No. RSA 174, II III (Holotype PREM, Isotype M).
- On Pteronia divaricata. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 24.7’’, E 18° 18’ 34.7’’, 19.1X.2002, leg. M. Mennicken No. RSA 189, II Il (Paratypes PREM, M).
- On Pteronia divaricata. Namibia, Bezirk Liideritz, 35 km N of Rosh Pinar (Route to Aus), 2-4 km west of the route, 29.X.1983, leg. B. Leuenberger, Th. Raus & Ch. Schiers No. 3248, II (B 10 9009304).
)
- On cf. Pteronia spec. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 24.5’’, E 18° 18’ 35.5’’, 19.1X.2002, leg. M. Mennicken No. RSA 194, II Ill (PREM, M).
The host genus Pteronia, which belongs to the Astereae, occurs with 70 species mainly in Namibia, Botswana, and South Africa. A few species are extending into Zimbabwe and into Australia (Herman, Retief, Koekemoer & Welman 2000).
As far as could be established, there is no known rust fungus either from the genus Pteronia or a similar known rust fungus from other genera of the family Asteraceae in southern Africa or in Australia. Therefore, the rust fungus is described as new.
ASTERACEAE - SONCHUS
Miyagia pseudosphaeria Jorst. (Jérstad 1961: 78).
Anamorph. Aecidium sonchi Johnst.
Syn. Puccinia pseudosphaeria Mont.
Syn. Puccinia sonchi Rob. in Desmaziéres 1849: 274. Lectotype on Sonchus arvensis, France, au- tumn.
Syn. Puccinia pseudo-sphaeria Mont.
Anamorph. Aecidium sonchi Westend. (Westendorp 1861: 649). Type on Sonchus oleraceus, Bel- gium, near Courtrai (HCB No. 1160).
Syn. Uromyces sonchi Oudem.
Syn. Uromyces puccinioides Fautrey & Rolland in Roumeguére 1893: 25. Type on Sonchus arven- sis, France, VIII.1892, leg. F. Fautrey.
Syn. Puccinia tagananensis Magnus 1901: 297. Type on Sonchus radiatus Ait. Teneriffa, near Tagana, VII.1900, leg. Bornmiiller.
Syn. Peristemma sonchi (Rob.) Syd. (Sydow 1921: 175).
Syn. Peristemma pseudosphaeria (Mont.) J@rst. (Jgrstad 1956a: 280).
Pycnia and telia not seen. Aecia unknown.
Uredinia amphigenous on leaves, subepidermal, separate or scattered in irregular groups, yellowish pustules small, up to 1 mm wide, long covered by the epidermis, opening through an apical pore, later gaping, pulverulent, spots surrounding pustule up to 2 mm wide, often discoloured violet, not clearly margined on the host leaf. Paraphy- ses numerous, peripherally surrounding the sori, cylindrical to subclavate, marginally thicker at the apex, pale brown at the base, chestnut-brown at the apex, thick-walled. Urediniospores borne singly on pedicels, ovoid, ellipsoid to oblong ellipsoid, 26—40 x 15-24 um, echinulate, spore wall about 2—5 um thick, hyaline, germ pores obscure, only visible in optical sections by the reduced thickness of the spore wall, scattered and probably numerous, without papillae.
Specimens examined:
- On Sonchus cf. oleraceus L. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 273 52.5’, E 18° 16’ 49.2’, 08.1X.2002, leg. M. Mennicken No. RSA 125, II (PREM, M).
- On Sonchus cf. oleraceus. South Africa, Western Cape Province, BIOTA-observatory at Riverlands, S 33° 29’ 27.7”, E 18° 35’ 25.9’, 19.XI.2001, leg. Markus Goker No. RSA 16, IT (PREM, M).
- On Sonchus cf. oleraceus. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27° 42.2”, E 18° 16’ 28.4’’, 18.1X.2002, leg. M. Mennicken No. RSA 173, II (PREM, M).
Sonchus cf. oleraceus is one of the earliest weeds introduced into South Africa from Europe, having been recorded at the Cape as early as 1685 (Bromilow 2001). As far as could be established, Miyagia pseudosphaeria is new to the rust flora of South Africa.
10
ASTERACEAE - TARCHONANTHUS Uredo tarchonanthi Mennicken & Oberw. sp. nov. (Fig. 6)
Pycnia, aecia et telia ignota. Uredinia abaxialia, subepidermalia, aurantiaca vel ochracea vel cremeo-alba, usque ad 0.2 mm diam., immersa, exposita, pulverulenta, paraphysibus peripheralibus cylindraceis, clavatis, capitatis vel irregularibus, incurvatis, septatis, dilute aureo-brunneis, usque ad 100 um longis, circa 7-19 um latis, pariete basali circa 0.5—1 pm crasso, apicali incrassato usque ad 4 um. Urediniosporae subgloboideae, ovoideae, ellip- soideae, pyriformes vel oblongae, 26-41 (-45) x 17-26 um, pariete echinulato, dilute aureo-fusco vel hyalino, 1—3 um crasso.
In foliis Tarchonanthi littoralis P.P.J. Herman (Asteraceae).
Fig. 6: Uredo tarchonanthi on Tarchonanthus littoralis. Paraphyses and urediniospores (Holotype RSA 220). Scale bar = 10 um.
Pycnia, aecia, and telia unknown.
Uredinia abaxial on leaves, scattered, subepidermal, freshly orange and conspicuous, bleaching in the herbarium to ochraceous or cream-white and very inconspicuous as a result of the cream-white, hairy tomentum covering the abaxial leaf surface, single sori strongly immersed, up to 0.2 mm diameter, exposed, pulverulent, surrounded by numer- ous, mostly incurved paraphyses, paraphyses cylindrical, clavate, or capitate, often ir- regularly in outline, 1— to 3—septate, up to circa 100 um long and circa 7-19 um wide near the apex, wall of the paraphyses about 0.5—1 um thick basally, up to 4 um thick near the apex, yellow-brown to hyaline. Urediniospores sessile, subgloboid, ovoid, el- lipsoid, pyriform or oblong, 26-41 (-45) x 17-26 um, grossly echinulate, echinulation closely meshed, spore wall uniformly about 1—3 um thick, very pale brownish-yellow to hyaline, germ pores obscure, occasionally visible in optical sections, scattered and probably numerous, without papillae.
On the leaves of Tarchonanthus littoralis (Asteraceae).
Etymology. Named after the host plant, Tarchonanthus littoralis.
Specimen examined:
- On Tarchonanthus littoralis. South Africa, Western Cape Province, Cape Peninsula National Park, BIOTA- observatory at Olifantsbos, 23.[X.2002, leg. M. Mennicken No. RSA 220, II (Holotype PREM, Isotype M). In- fected with the hyperparasite Eudarluca caricis (Biv.) O.E. Erikss.
11
The genus Tarchonanthus forms together with the genus Brachylaena the tribe Tarcho- nantheae and is restricted to Africa, Madagascar and associated islands (Herman, Re- tief, Koekemoer & Welman 2000). Doidge (1927) described the rust fungus Uredo brachylaenae on several Brachylaena species in Natal, Transvaal, and Cape Province in South Africa. Uredo brachylaenae resembles Uredo tarchonanthi in several details like the peripheral, septate paraphyses, and the obscure germ pores. It differs from Uredo tarchonanthi in having smaller spores (18—23.5 x 15-17 um) with a thinner- walled epispore (about | um), and in having “minutely and rather sparsely verruculose- echinulate” spore wall. We examined the type specimens of Uredo brachylaenae (Holotype PREM 2525, Paratype PREM 1821, and Paratype PREM 982). Although the urediniospores often have larger dimensions than stated in Doidge (1927), they do not reach the values of our measurements. Furthermore, the echinulate spore wall of Uredo brachylaenae is uniformly about 1—1.5 um thick, the sori are not immersed but su- perepidermally developed, and the arrangement of the paraphyses is more consolidated in the lower parts of the sorus. Therefore, Uredo tarchonanthi is described as new.
ASTERACEAE - URSINIA Puccinia ursiniae R.G. Shivas (Shivas 1991: 379). (Fig. 7)
Type on Ursinia anthemoides (L.) Poir. Western Australia, Burns Beach Road, Wanneroo,
20.X.1985, leg. R. G. Shivas (PERTH 791873). QOOOG
OEBGSGO89
Fig. 7: Puccinia ursiniae on Ursinia anthemoides. Uredinio- and teliospores (RSA 14). Scale bar = 10 um.
Pycnia and aecia unknown.
Uredinia mostly amphigenous on leaves, rarely on stems and on peduncles, scattered, single sori round to elliptic in outline, small, up to 1 mm diameter, cinnamon-brown, early exposed, pulverulent, surrounded by the torn epidermis, without leaf spots sur- rounding uredinia. Urediniospores subgloboid, pyriform, ellipsoid or obovoid, 27—38 x 21-26 um, echinulate, spores borne singly on pedicels, spore wall about 1.5—2.5 um thick at the sides, about 2-4 um thick at the apex, golden-brown to pale brown, germ pores inconspicuous, 3, equatorial, with hyaline papillae that are largely developed. Telia amphigenous on leaves, on stems, on peduncles and on involucral bracts, (partly) developing from the uredinia, scattered or confluent, single sori on the leaves and on the
12
involucral bracts round to ellipsoid, up to 1 mm diameter, on the stems and the peduncles ellipsoid to oblong, up to 2 mm long, early exposed, cushion-like, pulveru- lent, surrounded by the torn epidermis, without leaf spots surrounding telia. Teliospores bicellular, ellipsoid or cylindrical, rounded, slightly apiculate or flattened at the apex, rounded or attenuate at the base, not or slightly constricted at the septum, 37-63 x 24— 34 um, spore wall verrucous, with spaced cavities, about 1.5—3 ym thick at the sides, about 5—10 um thick at the apex, chestnut-brown, germ pore in the upper cell apical, in the lower cell just below the septum, pedicel persistent, up to 120 um long, thin- or thick-walled, in general not collapsing, hyaline, sometimes obliquely inserted.
Specimens examined:
- On Ursinia anthemoides. South Africa, Western Cape Province, BIOTA-observatory at Elandsberg, S 33° 26’ 05.8’, E 19° 02’ 18.5’’, 18.X1.2001, leg. M. Mennicken No. RSA 9, III] (PREM, M).
- On Ursinia anthemoides. South Africa, Western Cape Province, BIOTA-observatory at Elandsberg, S 33° 26’ 01.2”’, E 19° 02’ 09.8”’, 21.1X.2002, M. Mennicken No. RSA 209, I II] (PREM, M).
- On Ursinia anthemoides. South Africa, Western Cape Province, BIOTA-observatory at Riverlands, S 33° 29’ 19.1°’, E 18° 35’ 20.2’’, 19.XI.2001, leg. M. Mennicken No. RSA 14, I III (PREM, M).
- On Ursinia anthemoides. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 17.2”’, E 18° 18’ 46.5’’, 21.XI.2001, leg. M. Mennicken No. RSA 24, (ID) I (PREM, M).
The genus Ursinia, which belongs to the Anthemideae, occurs with 39 species in southern Africa and Ethiopia. It is widespread, and especially prominent in Northern Cape (Namaqualand), Western Cape, and the Eastern Cape (Herman et al. 2000). Urs- inia anthemoides has been introduced in Australia and “has become a weed of roadsides and waste places on a variety of soils extending throughout the south-western region of the State“‘ (Marchant et al. 1987 in Shivas 1991).
Our collections agree well with the diagnosis of Puccinia ursiniae given by Shivas (1991). The type collection is from Western Australia, but Shivas (1991) checked the possibility that Puccinia ursiniae was indigenous to South Africa. He found that eight of 26 herbarium specimens of Ursinia anthemoides received from South Africa were in- fected with Puccinia anthemoides. The oldest South African specimen of Ursinia an- themoides infected with Puccinia ursiniae was collected in the year 1885.
CHENOPODIACEAE - SALSOLA
Uredo sp. (Fig. 8)
Pycnia, aecia and telia unknown or not seen, respectively.
Uredinia amphigenous on the elongated leaves of spherical, small, up to 1,5 cm wide, gall-shaped concrescences, subepidermal, scattered, irregularly rupturing pustules up to 1 mm wide, roundish or elliptic, cinnamon-brown, without spots surrounding pustules, early exposed and surrounded by the torn epidermis. Urediniospores subgloboid, pyri- form, ellipsoid or terete, 24-32 (—34) x 19-24 (—26) um, echinulate, spores borne singly on pedicels, spore wall uniformly about 1.5—2.5 um thick, golden to pale chestnut- brown, germ pores 3 to 6, scattered, with hyaline papillae.
Specimens examined:
- On Salsola cf. zeyheri (Moq.) Bunge. South Africa, Western Cape Province, BIOTA-observatory at Moedver- loren 208, S 31° 27’ 42.3’’, E 18° 26’ 28.4’’, 18.1X.2002, leg. M. Mennicken No. RSA 170, II (PREM, M).
- On Salsola cf. zeyheri. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27’ 42.3’, E 18° 26’ 30.6’’, 18.1X.2002, leg. M. Mennicken No. RSA 182, II (PREM, M).
PS
Fig. 8: Uredo sp. on Salsola cf. zeyheri. Urediniospores (RSA 170). Scale bar = 10 um.
As far as could be established, there is no known South African rust fungus on Salsola. Because of the existence of several Uromyces species with similar urediniospore di- mensions on Chenopodiaceae (Sydow & Sydow 1910) our results are inconclusive.
FABACEAE - ASPALATHUS Uredo aspalathi Mennicken & Oberw. sp. nov. (Fig. 9)
Pycnia, aecia et telia ignota. Uredinia amphigena, subepidermalia, cinnamomea, usque ad | mm diam., mox nu- da, pulverulenta. Urediniosporae globoideae, pyriformes vel late ellipsoideae, 24-32 x 22-27 um, pariete echinulato, aureo-fusco, 2.5—3.5 um crasso, 4-6 poris germinationis dispersis.
In foliis Aspalathi laricifoliae Bergius (Fabaceae)
Pycnia, aecia, and telia unknown.
Uredinia amphigenous on leaves, subepidermal, forming small prominent pustules, up to 1 mm wide, roundish or elliptic, irregularly rupturing, cinnamon-brown, without spots surrounding pustules, clearly margined on the host leaves, surrounded by the torn epidermis. Urediniospores borne singly on short pedicels, globoid, pyriform or (broad- ly) ellipsoid, partially tending to be triangular to pentagonal with broadly rounded edg- es, 24-32 x 22-27 um, echinulate, echinulation coarsely and irregularly meshed, spore wall uniformly about 2.5—3.5 um thick, golden to dark golden-brown, germ pores con- spicuous, 4 to 6, scattered, with hyaline papillae.
On the leaves of Aspalathus laricifolia (Fabaceae).
Etymology. Named after the host plant, Aspalathus laricifolia.
Fig. 9: Uredo aspalathi on Aspalathus laricifolia. Urediniospores (Holotype RSA 2). Scale bar = 10 um.
Specimens examined:
- On Aspalathus laricifolia. South Africa, Western Cape Province, Cape Peninsula National Park, BIOTA-ob- servatory at Olifantsbos, S 34° 15’ 46.1’’, E 18° 23’ 36.6’, 17.XI.2001, leg. M. Mennicken No. RSA 2, II (Ho- lotype PREM, Isotype M).
- On Aspalathus laricifolia. South Africa, Western Cape Province, Cape Peninsula National Park, BIOTA-ob- servatory at Olifantsbos, S 34° 15’ 40.3’’, E 18° 23’ 35.5’’, 23.1X.2002, leg. M. Mennicken No. RSA 215, II (Paratypes PREM, M).
14
The genus Aspalathus, which belongs to the subfamily Papilionideae and the tribe Cro- talarieae is with + 278 species the largest genus of flowering plants endemic in southern Africa (Germishuizen 2000). The center of distribution is in the Western Cape but ex- tends into KwaZulu-Natal.
As far as could be established, only microcyclic rust fungi have been described on Aspalathus so far: Uromyces bolusii Massee 1901: 168 (Type on Aspalathus pachyloba Benth., South Africa, Montagu Bath, leg. Bolus No. 7597), and Uromyces ventosa (as Teleutospora ventosa) Syd. (Sydow 1924: 235) (Type on Aspalathus sp. (as Borbonia sp.), South Africa, Stellenbosch, VUI.1923, leg. A. V. Duthie No. 1246).
FABACEAE - LEBECKIA
Aecidium dielsii Henn. (Hennings 1904: 34). (Fig. 10)
Type on Crotalaria sp. South Africa, Clanwillian, 1900, leg. Diels No. 1145 (B).
Pycnia unknown.
Aecia aecidioid, amphigenous on leaves, on flowers, and on stems, covering the whole stem surface over several centimeters, often causing hypertrophy of the stems, without spots surrounding aecia, aecial cups reduced to salient, erumpent pustules, circa 400-500 um diameter, spore mass orange, yellow to cream-white, peridium exserted, slit right to the base in narrow segments, yellowish to cream-white, up to 2.2 cm long, cells of the peridium medium connected, oblong, outer wall striate, circa 3—7 um wide, inner wall inconspicuously striate, circa 1.5—3 um wide. Aeciospores angular globoid, subgloboid, ellipsoid to oblong, 22—33 x 18—26 um, spore wall about 2-4 um thick, up to 5.5 um thick in one edge, finely verrucous, yellowish to hyaline, germ pores normally invisible, partly visible in optical sections, probably scattered and numerous, without
= nOo0d
Fig. 10: Aecidium dielsii on Lebeckia sp. Peridium cells and aeciospores (RSA 185). Scale bar = 10 um.
Specimen examined:
- On Lebeckia sp. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 32.9’’, E 18° 18’ 23.1’’, 19.1X.2002, leg. M. Mennicken No. RSA 185, I (PREM, M).
Our collection agrees well with the diagnosis of Aecidium dielsii. As far as could be established, Aecidium dielsii is known only from the type collection on Crotalaria sp. Thus, Lebeckia could be proven as a new host genus. Both host genera belong to the tribe Crotalarieae in Papilionoideae (Germishuizen 2000).
15
FABACEAE - PSORALEA
cf. Phakopsora pachyrhizae Syd. & P. Syd. (Sydow & Sydow 1914: 108).
Type on Pachyrhizus angulatus Rich. Formosa, Taihoku, 25.XII.1913, leg. Y. Fujikuro No. 37. For synonyms and figures see Ono, Buritica & Hennen (1992).
Pycnia and aecia unknown. Telia not seen.
Uredinia amphigenous on leaves, subepidermal, separate or scattered, pustules small, up to 0.25 mm diameter, roundish, pale brown, opening through a central aperture, pul- verulent, not clearly margined on the host leaves, spots surrounding pustule up to 1 cm wide, roundish or irregularly in outline, discoloured brown to dark brown, not clearly margined at the host leaves, single sori surrounded by paraphyses arising from peridioid pseudoparenchyma, also with hymenial paraphyses, paraphyses cylindrical to clavate, up to 45 um long and up to 12 um wide apically, wall of the paraphyses thin at the sides, up to 11 um thick at the apex, pale golden to hyaline. Urediniospores subgloboid, obo- void, pyriform to ellipsoid, 20—28 (—32) x 13-18 um, echinulate, spore wall uniformly about | um thick, pale yellowish-brown to hyaline, germ pores rather obscure, probably 2 to 8, scattered or + equatorial, without or with hyaline papillae that are very weakly developed.
Specimen examined:
- On Psoralea pinnata L. South Africa, Western Cape Province, Kirstenbosch Botanical Garden, 29.1X.2002, leg. M. Mennicken No. RSA 221, II (PREM, M).
According to Ono, Buritica & Hennen (1992), Phakopsora pachyrhizi infects a wide range of Leguminosae in Austro-Asia and Africa. Without teliospores Phakopsora pachyrhizi can not be delimited from Phakopsora meibomiae (Arthur) Arthur 1917: 509 (Syn. Phakopsora psoraleae Jackson 1931: 346. Type on Psoralea glandulosa L. Bo- livia, Sorata). Phakopsora meibomiae attacks several Leguminosae in the New World. Because of the geographic origin of our collection from South Africa, we refer to this rust fungus as Phakopsora pachyrhizae. As far as could be established Psoralea seems to be a new host genus for Phakopsora pachyrhizi.
GERANIACEAE - PELARGONIUM
Puccinia granularis Kalchbr. & Cooke in Kalchbrenner 1882: 22.
Type on Pelargonium sp. South Africa, P. Natal, No. 10.
Anamorph. Aecidium pelargonii Thiim. (Thiimen 1877: 411). Type on Pelargonium alchemil- loides Willd. South Africa, Boschberg near Somerset-East, Promont, XI.1876, leg. MacOwan. Anamorph. Uredo pelargonii Thiim. (Thiimen 1878: 355). Type on Pelargonium alchemilloides. South Africa, Boschberg near Somerset-East, Promont, XI.1876, leg. MacOwan.
Syn. Puccinia pelargonii (Thiim.) P. Syd. & Syd. (Sydow & Sydow 1904: 469).
Pycnia, aecia, and telia not seen.
Uredinia abaxial on leaves, separate, scattered or in irregularly crowded groups, red- dish-brown, forming round or irregular shaped sori on indefinite leaf spots, early exposed, pulverulent, surrounded by the torn epidermis. Urediniospores globoid, sub- globoid, or ellipsoid, 24-29 x 21-24 um, echinulate, spores borne singly on pedicels, spore wall uniformly about 2.5—3.5 um thick, golden-brown, germ pores 2 (to 3), equa- torial, without papillae.
16
Specimen examined:
- On Pelargonium grossularioides (L.) L’Hér. South Africa, Western Cape Province, BIOTA-observatory at Elandsberg, S 33° 26’ 03.2”’, E 19° 02’ 23.5’’, 18.X1.2001, leg. M. Mennicken No. RSA 10, II (PREM, M).
Our collection agrees well with the description of Puccinia granularis given by Doidge (1927). In South Africa one collection of Puccinia granularis on Pelargonium grossu- larioides is known from the Western Cape Province, University of Cape Town grounds, 31.1V.1934, leg. J.P.H. Acocks No. 2443 (Jgrstad 1956b). This collection is not men- tioned in Crous, Philip & Baxter (2000).
LAMIACEAE - MENTHA
Puccinia menthae Pers.
For synonyms see Sydow & Sydow (1904).
Pycnia, aecia, and telia not seen.
Uredinia abaxial on leaves, on stems, subepidermal, pale cinnamon-brown, separate, scattered, or arranged in irregular or concentric groups, single sori roundish, ellipsoid, or irregularly in outline, up to 1 mm diameter, first covered by the epidermis, early ex- posed, pulverulent, surrounded by the torn epidermis, mostly with yellowed, brightened or browned leaf spots surrounding uredinia. Urediniospores globoid, subgloboid, or el- lipsoid, 22—29 x 18-22 um, finely echinulate, spores borne singly on pedicels, spore wall uniformly about 1—2 um thick, pale golden-brown to hyaline, germ pores incon- spicuous, 2 to 3, + equatorial, occasionally one additional apical germ pore, with hyaline papillae that are weakly developed.
Specimen examined:
- On Mentha cf. longifolia (L.) L. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 53.8’’, E 18° 16’ 51.5’’, 08.1X%.2002, leg. M. Mennicken No. RSA 127, IJ (PREM, M).
Puccinia menthae, which is known from the provinces Gauteng and KwaZulu-Natal in South Africa (Crous, Phillips & Baxter 2000), seems to be new to the rust flora of Northern Cape.
ORCHIDACEAE - BARTHOLINA
Uromyces silksvleyensis Mennicken & Oberw. sp. nov. (Fig. 11)
Pycnia, aecia et uredinia ignota. Telia in foliis, plurimum abaxialia, raro adaxialia vel in petiolis, subepidermalia, nigro-brunnea, usque ad 1.2 mm diam., epidermide diu tecta, pulverulenta. Teliosporae unicellulares, globoide- ae, subgloboideae vel ellipsoideae, apice rotundato, leviter acuto vel leviter applanato, base rotundata vei leviter attenuata, 26-36 x 23-29 (—33) um, pariete laevigato, castaneo, lateraliter 0.5—1.5 um crasso, apicaliter 3-4 (— 5) um crasso, poris germinationis inconspicuis, pedicello dilute aureo vel hyalino, usque ad 35 um longo.
In foliis cf. Bartholinae burmannianae Ker-Gawl. (Orchidaceae).
Pycnia, aecia, and uredinia unknown.
Telia abaxial on leaves, occasionally adaxial, on petioles, scattered in confluent groups which can cover the whole leaf surface, subepidermal, single sori roundish, ellipsoid or irregularly in outline, up to 1.2 mm diameter, dark chocolate-brown to black, long covered by the epidermis which ruptures lately but always partly veiled, compact, pul- verulent, without leaf spots surrounding telia. Teliospores unicellular, globoid, sub- globoid to ellipsoid, often somewhat angular and irregularly, occasionally broader than long, rounded, slightly flattened, or slightly acute at the apex, rounded or slightly atten-
Li
uate at the base, 26-36 x 23-29 (—33) um, spore wall smooth, about 0.5—1.5 um thick at the sides, about 3-4 (—5) um thick at the apex, becoming progressively thicker from the base towards the apex, widest thickness not inevitably in the opposite of the pedicel, often obliquely displaced, chestnut-brown, germ pore obscure, probably apical, pedicel persistent, up to 35 um long, thin-walled, collapsing, yellowish to hyaline, sometimes obliquely inserted.
On the leaves and petioles of cf. Bartholina burmanniana (Orchidaceae).
Etymology. Named after the neighbouring field name, Silksvley.
VO0994
Fig. 11: Uromyces silksvleyensis on cf. Bartholina burmanniana. Teliospores (Holotype RSA 218). Scale bar = 10 um.
Specimen examined:
- On cf. Bartholina burmanniana. South Africa, Western Cape Province, Cape Peninsula National Park, BIOTA- observatory at Olifantsbos, S 34° 15’ 54.3’’, E 18° 24 03.2’’, 23.1X.2002, leg. M. Mennicken No. RSA 218, III (Holotype PREM, Isotype M).
As far as could be established, there is neither a known Uromyces species on Orchidace- ae in southern Africa nor is there any known Uromyces species on the host genus Bar- tholina, which, with two species, is endemic to southern Namibia, and to Northern, to Western and to Eastern Cape (Kurzweil 2000). Therefore, the rust fungus is described as new.
POACEAE - BROMUS
Puccinia hordei G.H. Otth.
For synonyms and figures see Cummins (1971).
Pycnia and aecia not seen.
Uredinia mostly adaxial on leaves, creamish-white to ochraceous, subepidermal, form- ing long streaks between the nerves of the leaves, early exposed, pulverulent, surround- ed by the torn epidermis. Urediniospores globoid, subgloboid, ovoid, pyriform or ellipsoid, 21-33 (—39) x 19-29 um, finely echinulate, spores borne singly on pedicels, spore wall uniformly about 1—2.5 um thick, pale yellowish, pale brownish to hyaline, germ pores inconspicuous, 10 to 15, scattered, with hyaline papillae that are weakly de- veloped.
Telia amphigenous on leaves, mostly abaxial, and on sheaths, dark chocolate-brown to blackish, forming long, confluent streaks between the nerves of the leaves, long remaining covered by the epidermis which later longitudinally ruptures, loculate by brown, palisade-wise paraphyses, compact. Teliospores bicellular, variable in shape and size, ellipsoid, clavate, oblong or irregularly in outline, often angular, rounded,
18
acuminate or flattened at the apex, rounded or attenuate at the base, slightly constricted at the septum, (30—) 36-63 x 17-32 (—36) um, spore wall smooth or with longitudinal surface ridges, about 1—2 um thick at the sides, about 3—9 um thick at the apex, yellow- brown to chestnut-brown, mostly paler towards the base, often paler towards the apex, germ pores obscure, pedicel persistent, up to 20 um long, often truncate immediately below the attachment, thin-walled, collapsing, yellowish to hyaline, often obliquely in- serted. Mesospores and tricellular spores existing.
Specimens examined:
- On Bromus pectinatus Thunb. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 47.5”, E 18° 16’ 45.3’’, 27.X1.2001, leg. M. Mennicken No. RSA 67, IJ HI (PREM, M).
- On Bromus pectinatus. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’? 30.8’, E 17° 33’ 11.9’, 11.1X.2002, leg. M. Mennicken No. RSA 137, II III (PREM, M).
- On Bromus pectinatus. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 31.0’, E 17° 32’ 55.7’’, 11.1X.2002, leg. M. Mennicken No. RSA 142, II If[ (PREM, M).
- On Bromus pectinatus. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 49.3”, E 18° 16° 43.4’, 27.X1.2001, leg. M. Mennicken No. RSA 69, II Ill (PREM, M).
- On Bromus pectinatus. South Africa, Western Cape Province, BIOTA-observatory at Flaminkvlakte 111, S 31° 17° 07.1’, E 18° 35’ 46.6’’, 06.[X.2002, leg. M. Mennicken No. RSA 116, II (PREM, M).
- On Bromus pectinatus. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27 48.0’’, E 18° 26’ 31.4’’, 05.1X.2002, leg. M. Mennicken No. RSA 113, Il UI (PREM, M).
- On Bromus pectinatus. South Africa, Western Cape Province, BIOTA-observatory at Luiperskop 211, S 31° 17° 37.1’, E 18° 36’ 14.6’’, 07.1X.2002, leg. M. Mennicken No. RSA 118, II II (PREM, M).
- On Bromus pectinatus. South Africa, Western Cape Province, BIOTA-observatory at Luiperskop 211, S 31° 17’ 30.7’, E 18° 36’ 04.0”’, 23.X1.2001, leg. M. Mennicken No. RSA 35, III (PREM, M).
- On Bromus pectinatus. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 28.8’, E 18° 18’ 29.9’, 19.1X.2002, leg. M. Mennicken No. RSA 201, IJ II (PREM, M).
Puccinia hordei agrees well with the description given in Cummins (1971). As far as could be established, there is only one collection of Puccinia hordei (on Bromus pecti- natus) in South Africa (Gjzrum 1988). This collection is not mentioned in Crous, Phil- lips & Baxter (2000). In South Africa the host plant Bromus pectinatus is regarded as an exotic plant from Eurasia (Bromilow 2001).
POACEAE - CHAETOBROMUS
Uromyces chaetobromi Gjerum 1988: 371. (Fig. 12)
Type on Chaetobromus schraderi Stapf. South Africa, Cape Province, Clanwillian distr. (as Chan- willian distr.), 07.XI.1974, leg. Van Breda No. 4302 (Holotype K, Isotype NPPI).
Pycnia and aecia unknown.
Uredinia amphigenous on leaves, subepidermal, cinnamon-brown, forming small ellip- tic to oblong, later confluent patches between the nerves of the leaves, early exposed, pulverulent, surrounded by the torn epidermis. Urediniospores globoid, subgloboid or ellipsoid, 24-33 x 22-31 um, echinulate, spores borne singly on pedicels, spore wall uniformly about 2—2.5 (—3) um thick, yellow-brown, germ pores 6 to 12, scattered, with hyaline papillae that are weakly developed.
Telia amphigenous on leaves, subepidermal, blackish, (partly) developing from the uredinia, forming small elliptic to oblong patches, long covered by the epidermis, later- exposed, compact to cushion-like, surrounded by the torn epidermis. Teliospores uni- cellular, 21-31 x 18-27 um, irregularly in shape, mostly angular globoid, subgloboid to ellipsoid, sometimes broader than long, rounded, acute or flattened at the apex, rounded
A=)
or attenuate at the base, chestnut-brown, spore wall smooth, about 1—2 um thick at the sides, about 3—7 um thick at the apex, germ pore apical, pedicel persistent, up to 55 um long, thin-walled, collapsing, yellow-brownish to hyaline, often obliquely inserted.
Fig. 12: Uromyces chaetobromi. Uredinio- and teliospores on Chaetobromus dregeanus (RSA 135). Scale bar = 10 um.
Specimens examined:
- On Chaetobromus dregeanus Nees. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 53.9’’, E 18° 16’ 49.7°’, 08.1X.2002, leg. M. Mennicken No. RSA 126, II (PREM, M).
- On Chaetobromus dregeanus. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 31.0’, E 17° 32’ 45.4’, 11.1X.2002, leg. M. Mennicken No. RSA 135, II II] (PREM, M).
- On Chaetobromus dregeanus. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27’ 40.5’’, E 18° 26’ 30.5’’, 05.1X.2002, leg. M. Mennicken No. RSA 102, II II] (PREM, M).
Our collections agree well with the characteristics of Uromyces chaetobromi given in the diagnosis (Gjzrum 1988). No uredinia but only urediniospores were described from the type collection. The number of germ pores of the urediniospores is wider in our col- lections (6 to 12) compared to the type collection (7 to 8). As far as could be established, Uromyces chaetobromi is only known from the type collection. The type collection is not mentioned in Crous, Phillips & Baxter (2000). Chaetobromus dregeanus seems to be a new host plant.
POACEAE - EHRHARTA
Uromyces ehrhartae-giganteae Doidge 1927: 207 emend. (Fig. 13)
Type on Ehrharta thunbergii Gibbs Russell (= E. gigantea Thunb.), South Africa, Mobray, Cape Province, 10.11.1914, leg. Van der Merve No. 7392.
Anamorph. Uredo ehrhartae-calycinae Doidge 1948: 907. Type on Ehrharta calycina J.E. Sm. South Africa, Stellenbosch, leg. Verwoed, Herb. Stell. Elsenburg Coll. Agric. 41 (34098). Pycnia and aecia unknown.
Uredinia amphigenous on leaves, mostly adaxial, cinnamon-brown, forming long and narrow, later confluent streaks between the nerves of the leaves, early exposed, pulver- ulent, surrounded by the torn epidermis. Urediniospores globoid, pyriform or broadly obovoid, 20-31 x 19-26 um, echinulate, spores borne singly on pedicels, spore wall uniformly about 2.5—4 um thick, golden to chestnut-brown, germ pores 6 to 8, scattered, with inconspicuous, hyaline papillae that are weakly to broadly developed.
20
Telia mostly adaxial on leaves, rarely abaxial, on sheaths or stems, dark chocolate- brown, forming patches and long, confluent streaks, exposed, cushion-like, surrounded by the torn epidermis. Teliospores unicellular, 26-43 (-46) x 19-32 um, tending to be dimorphic with broadly ellipsoid or globoid spores that are mostly more than 25 um broad and ellipsoid or oblong-ellipsoid spores that are mostly less than 25 um broad, occasionally asymmetrically angular, golden-brown to chestnut-brown, rounded, flattened or acute at the apex, rounded or attenuate at the base, spore wall smooth, about 1-4 um thick at the sides, up to 12 um thick at the apex, germ pore apical, pedicel per- sistent, up to 170 um long, not collapsing, yellow-brownish to hyaline, thick-walled, pedicel wall about 1.5—3 um thick.
Fig. 13: Uromyces ehrhartae-giganteae. Uredinio- and teliospores on Ehrharta villosa (RSA 190). Scale bar = 10 um.
Specimens examined:
- On Ehrharta calycina. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 26.6’’, E 18° 18’ 32.0’, 19.1X.2002, leg. M. Mennicken No. RSA 192, II (PREM, M).
- On Ehrharta villosa Schult. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 33.8’, E 18° 18’ 22.4’, 19. IX.2002, leg. M. Mennicken No. RSA 190, II III (PREM, M).
- On Ehrharta villosa. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 15.6’, E 18° 18’ 37.6’’, 21.X1.2001, leg. M. Mennicken No. RSA 20, III (PREM, M).
The urediniospores of our collections agree well with the specifications of Uredo ehrhartae-calycinae, which are given in the diagnosis and revised by Cummins (1971): (20—) 22-27 (—29) um x (17—) 19-21 um and (4 to) 5 to 8 scattered germ pores. Only the maximum broadness of the urediniospores is 5 um more in our collections.
Our teliospores coincide with the dimorphic characteristic of Uromyces ehrhartae-gi- ganteae, but they reach a larger size than in the diagnosis (Doidge 1927) or in the de- scription of Cummins (1971), respectively. Cummins (1971) scaled up the size of the teliospores given in the diagnosis and specifies for the type: “spores (23—) 26-36 (—38; 40) x (16—) 19-24 (—26) um”.
Summarizing, the deviation in the maximum broadness of the urediniospores and in the size of the teliospores does not justify a new species, all the more so, as the habitats are close-by, and the teliospores were found on a host plant different from the host of the type collection. We assume that our collections belong to Uromyces ehrhartae-gigante- ae, which is synonymous with Uredo ehrhartae-calycinae. Ehrharta villosa seems to be a new host plant for this rust fungus.
pA |
Uromyces quaggafonteinus Mennicken & Oberw. sp. nov. (Fig. 14)
Pycnia et aecia ignota. Uredinia amphigena, subepidermalia, vulpina, mox nuda, pulverulenta. Urediniosporae globoideae vel late ellipsoideae, 26-36 x 24-32 um, pariete echinulato, luteo-fusco vel dilute castaneo, 2.5-4 um crasso, 5—9 poris germinationis dispersis. Telia amphigena, subepidermalia, nigro-brunnea, mox nuda, pulv- inata. Teliosporae unicellulares, globoideae vel subellipsoideae, apice rotundato, leviter acuto vel leviter applan- ato, base rotundata, 28-39 x 24-32 um, pariete laevigato, castaneo, lateraliter 24 um crasso, apicaliter usque ad 12 um crasso, poro germinationis apicali, pedicello dilute aureo-fusco vel hyalino, usque ad 140 um longo.
In foliis Ehrhartae calycinae J.E. Sm. (Poaceae).
TTT e SY
Fig. 14: Uromyces quaggafonteinus. Uredinio- and teliospores on Ehrharta calycina (Paratype RSA 49). Scale bar = 10 um.
Pycnia and aecia unknown.
Uredinia amphigenous on leaves, subepidermal, foxy red, forming small, elliptic to ob- long, later confluent patches between the nerves of the leaves, early exposed, pulveru- lent, surrounded by the torn epidermis. Urediniospores globoid or broadly ellipsoid, golden to pale chestnut-brown, 26-36 x 24—32 um, echinulate, borne singly on pedicels, spore wall uniformly about 2.5—4 um thick, germ pores 5 to 9, scattered, with incon- spicuous, hyaline or yellowish papillae that are weakly developed.
Telia amphigenous on leaves, subepidermal, dark chocolate-brown, forming small, elliptic to oblong, later confluent patches, between the nerves of the leaves, exposed, cushion-like, surrounded by the torn epidermis. Teliospores unicellular, globoid to su- belliptic, rounded, slightly flattened or slightly acute at the apex, rounded at the base, 28-39 x 24-32 um, chestnut-brown, spore wall smooth, about 2—4 um thick at the sides, up to 12 um thick at the apex, germ pore apical, pedicel persistent, up to 140 um long, not collapsing, brownish-yellow to hyaline, thick-walled, pedicel wall up to 3 um thick, occasionally obliquely inserted.
On the leaves of Ehrharta calycina. (Poaceae)
Etymology. Named after the collection site, Quaggafontein.
Specimens examined:
- On Ehrharta calycina. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 31.1°’, E.17° 32’ 55.6’’, 11.1X.2002, leg. M. Mennicken No. RSA 139, II II (Holotype PREM, Isotype M).
De.
- On Ehrharta calycina. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 25.9’, E 17° 32’ 53.0’, 26.X1.2001, leg. M. Mennicken No. RSA 49, II III (Paratypes PREM, M).
Uromyces quaggafonteinus differs from Uromyces ehrhartae-giganteae (see above) in the lack of oblong-ellipsoid or asymmetrical angular teliospores. The mean length of the teliospores is shorter, the mean broadness of the teliospores is wider, the echinulation of the urediniospores is coarser and the size of the urediniospores is bigger than in Uro- myces ehrhartae-giganteae. Only with teliospores a precise determination of these two rust fungi is not possible.
POACEAE - HOLCUS, KARROOCHLOA, SCHISMUS, TRIBOLIUM
Uromyces holci J¢érst. (Jgrstad 1956b: 577). (Fig. 15)
Type on Holcus setiger Nees, South Africa, Northern Cape Province, between Pedroskloff and Leliefontein, leg. Drege (S).
Syn. Uromyces schismi J¢rst. (Jgrstad 1956b: 577). Type on Schismus scaberrimus Nees, locality unknown “but the collection is obviously from S. Africa, where the host belongs”.
i
Fig. 15: Uromyces holci. Uredinio- and teliospores. Scale bar = 10 um.
7 @)
Pycnia and aecia unknown.
Uredinia adaxial on leaves, foxy red, forming patches and stripes between the nerves of the leaves, early exposed, surrounded by the torn epidermis. Urediniospores globoid, broadly ovoid, or broadly ellipsoid, golden to pale chestnut-brown, 21—31 (—34) x 18-28 um, finely echinulate, spores borne singly on pedicels, spore wall uniformly about 1.5—2.5 (—3) um thick, germ pores 7 to 11, scattered, with hyaline papillae. Telia adaxial on leaves, blackish, forming small, later confluent patches, early exposed, cushion-like, surrounded by and arched over by the torn epidermis. Teliospores unicel- lular, variable in shape and size, 19-33 x 15-29 um, dimorphic with broadly ellipsoid or globoid spores tending to be chestnut-brown and with globoid, ellipsoid or oblong- ellipsoid, asymmetrically angular, variable spores tending to be golden to pale chestnut- brown, spore wall smooth, about 1—3 um thick at the sides, about 2—8 um thick at the apex, germ pore apical, pedicel persistent, up to 80 um long, brownish-yellow, thick- walled and predominantly not collapsing in the chestnut-brown spores, thin-walled and collapsing in the golden to pale chestnut-brown spores.
25
Specimens examined:
- On cf. Holcus setiger Nees. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Reserve, S 32° 36’ 17.4’’, E 18° 18’ 45.3’, 21.X1.2001, leg. M. Mennicken No. RSA 23, II III (PREM, M).
- On cf. Holcus setiger. South Africa, Western Cape Province, BIOTA-observatory at Rocherpan Nature Re- serve, S 32° 36’ 32.5’’, E 18° 18’ 38.6’’, 21.X1.2001, leg. M. Gdker No. RSA 26, II Ill (PREM, M).
- On Karroochloa cf. schismoides (Stapf ex Conert) Conert & Tuerpe. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 23.0’, E 17° 33’ 06.2’’, 26.X1.2001, leg. M. Mennicken No. RSA 51, I Tl (PREM, M).
- On Schismus barbatus (Loefl. ex L.) Thell. Namibia, Liideritz-Siid, Farm Spitzkop LUS, 2716 DC, 25.1X.1977, leg. H. Merxmiiller & W. Giess, IJ II] (WIND 46700).
- On Schismus barbatus. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 49.7’, E 18° 16’ 44.2’, 27.X1.2001, leg. M. Mennicken No. RSA 66, II (PREM, M).
- On Schismus barbatus. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 22.8’’, E 17° 33’ 13.0’’, 26.X1.2001, leg. M. Mennicken No. RSA 50, II Il (PREM, M).
- On Schismus barbatus. South Africa, Northern Cape Province, BIOTA-observatory at Leliefontein 624, S 30° 23’ 36.2”’, E 18° 16’ 35.8’, 27.X1.2001, leg. M. Mennicken No. RSA 57, II Il (PREM, M).
- On Schismus barbatus. South Africa, Northern Cape Province, BIOTA-observatory at Leliefontein 624, S 30° 23’ 36.4’, E 18° 16’ 37.6’, 27.X1.2001, leg. M. Mennicken No. RSA 58, II III (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Flaminkvlakte 111, S 31° 17’ 05.1’, E 18° 35’ 48.8”’, 23.X1.2001, leg. M. Mennicken No. RSA 30, II II] (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Luiperskop 211, S 31° 17 35.4’’, E 18° 36’ 04.0’’, 23.X1.2001, leg. M. Mennicken No. RSA 36, I Ill (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 27 40.1", E 18° 26’ 31.6’, 05.1X.2002, leg. M. Mennicken No. RSA 103, I II (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Luiperskop 211, S 31° 17° 37.1", E 18° 36’ 14.6’, 07.1X.2002, leg. M. Mennicken No. RSA 119, II Il (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Luiperskop 211, S 31° 17° 13.7’, E 18° 36’ 14.2’, 07.1X.2002, leg. M. Mennicken No. RSA 120, II Ill (PREM, M).
- On Schismus barbatus. South Africa, Western Cape Province, BIOTA-observatory at Moedverloren 208, S 31° 28’ 00.7’, E 18° 26’ 25.6’’, 26.X1.2001, leg. M. Mennicken No. RSA 104, III (PREM, M).
- On Tribolium echinatum (Thunb.) Renvoize (= Lasiochloa echinata (Thunb.) Adamson), South Africa, North- ern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 31.0’, E 17° 32’ 35.6’, 11.1X.2002, leg. M. Mennicken No. RSA 141, II III (PREM, M).
- On Tribolium echinatum . South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 29.4’, E 17° 33’ 05.7’, 11.1X.2002, leg. M. Mennicken No. RSA 144, II If (PREM, M).
Our collections agree well with the range of spore dimensions given in the diagnoses of Uromyces holci and Uromyces schismi. Dimensions range from 23-29 x 20—24 um for the urediniospores, and from 17-32 x 15-25 um for the teliospores. The number of germ pores in the urediniospores of our collection is 7 to 11, noticeably higher than the number given in the diagnoses, where 6 or 7, respectively, scattered germ pores are mentioned. Cummins (1971) listed the number of germ pores as varying from 7 to 9. In his figure one urediniospore has 9 and one urediniospore has 11 germ pores. We hypo- thesise that Uromyces holci has 7 to 11 germ pores in its urediniospores.
As far as could be established, Uromyces holci is new to the rust flora of Namibia. Kar- roochloa schismoides, Schismus barbatus and Tribolium echinatum seem to be new host plants. Uromyces holci is the first known rust fungus that infects the grass genera Karroochloa and Tribolium. Whilst the host genus Holcus belongs to the subfamily Pooideae and the tribe Aveneae, the three host genera Karroochloa, Schismus, and Tri- bolium belong to the subfamily Arundinoideae and the tribe Arundineae (Fish 2000).
POACEAE - Polypogon
Puccinia polypogonis Speg. (Spegazzini 1909: 300). (Fig. 16)
Type on Polypogon monspeliensis, Argentina, near Lake Muster, Patagonia, XII.1902 (LPS, Iso- type PUR).
24
Anamorph. Uredo polypogonis Speg. (Spegazzini 1899: 240). Lectotype on Polypogon monspe- liensis (L.) Desf. Argentina, Puerto Deseado, 1885/1886.
Pycnia and aecia not known.
Uredinia amphigenous on leaves, mostly adaxial, and on sheaths, foxy red, subepider- mal, forming partially confluent stripes between the nerves of the leaves, early exposed, pulverulent, surrounded by the torn epidermis. Urediniospores globoid to broadly el- lipsoid, golden-brown, 22—29 x 19-27 um, echinulate, spores borne singly on pedicels, spore wall uniformly about 1.5—3 um thick, germ pores 6 to 8, with hyaline papillae that are largely developed.
Telia amphigenous on leaves, blackish-brown, subepidermal, (partly) developing from the uredinia, forming small, ellipsoid to oblong patches between the nerves of the leaves, early exposed, compact to cushion-like, surrounded by the torn epidermis. Teliospores in general bicellular, 31—51 x 15—25 (—27) um, irregularly in shape and size, often oblique, varying from clavate to ellipsoid, occasionally diorchidioid, rounded, apiculate or flattened at the apex, attenuate or rounded at the base, constricted at the sep- tum, brown, spore wall smooth, about 1.5—2.5 um thick at the sides, about 2 to 9 um thick at the apex, germ pores apical and just below the septum, pedicel persistent, up to 55 um long, mostly collapsing, often obliquely inserted, brownish, just below the sep- tum of the lower cell darker tinted. Mesospores commonly developed.
JI0¢
Fig. 16: Puccinia polypogonis. Uredinio- and teliospores on Polypogon monspeliensis (WIND 41741, WIND 63839). Scale bar = 10 um.
Specimens examined:
- On Polypogon monspeliensis. Namibia, Liideritz — Siid, Loreleifelsen- und Kupfermine und Umgebung, im Flussbett des Oranje, 2816 BB Oranjemund, 02.X.1977, leg. H. Merxmiiller & W. Giess No. 32454, Il (WIND 41741).
- On Polypogon monspeliensis. Namibia, Nuop Rivier, 3 km noérdlich Einmiindung in den Oranje, 2817 AA Vioolsdrif, 02.X.1975, leg. W. Giess No. 13837, (ID) II] (WIND 41744).
- On Polypogon monspeliensis. Namibia, Kaokoland Area 2, Skeleton Coast Park, Uniab River, 2013 AA, 50 m, 22.X.1988, leg. C. J. Ward & J. D Ward No. 10480, (I) HI (WIND 63839).
- On Polypogon monspeliensis. South Africa, Northern Cape Province, BIOTA-observatory at Leliefontein 624, S 30° 23’ 39.5’’, E 18° 16’ 40.9’’, 27.X1.2001, leg. M. Mennicken No. RSA 60, II II (PREM, M).
25
- On Polypogon monspeliensis. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 49.1’, E 18° 16’ 45.2”’, 27.X1.2001, leg. M. Mennicken No. RSA 68, II II (PREM, M).
- On Polypogon monspeliensis. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 50.4’’, E 18° 16° 45.8’’, 27.X1.2001, leg. M. Mennicken No. RSA 70, II II (PREM, M).
- On Polypogon monspeliensis. South Africa, Northern Cape Province, BIOTA-observatory at Remhoogte 416, S 30° 23’ 43.0’’, E 18° 16’ 43.7’, 27.X1.2001, leg. M.Goker No. RSA 74, II (PREM, M).
- On Polypogon monspeliensis. South Africa, Northern Cape Province, BIOTA-observatory at Quaggafontein 478, S 30° 11’ 18.4’’, E 17° 33’ 01.2’’, 26.XI.2001, leg. M. Mennicken No. RSA 55, Il (PREM, M). This col- lection is double infected with Puccinia coronata Corda (only teliospores).
- On Polypogon monspeliensis. South Africa, Western Cape Province, BIOTA-observatory at Elandsberg, S 33° 25’ 55.4’’, E 19° 02’ 14.7’’, 18.X1.2001, leg. M. Mennicken No. RSA 7, II (PREM, M).
As far as could be established, Puccinia polypogonis is new to the rust flora of Namibia. A South African collection from Saron in the Cape Province by Schlechter in 1890 (Jgrstad 1956) is not cited in Crous, Phillips & Baxter (2000). The host plant Polypogon monspeliensis was introduced from Europe and Asia and is now naturalised in many parts of South Africa, especially in coastal areas (Bromilow 2001).
SANTALACEAE - THESIUM
Puccinia cf. stonemaniae Syd., P. Syd & Pole-Evans in Sydow & Sydow 1912: 437. (Fig. 17)
Type on Thesium sp. South Africa, Bain’s Kloof near Wellington, 21.11.1912, leg. E. M. Doidge.
Anamorph. Aecidium osyridicarpi Massee 1911: 225. Type on Osyridicarpos natalensis DC. South Africa, Natal, Tabamhlope, leg. Wood No. 527.
Syn. Puccinia pulvinata Massee 1911: 224 (nec. Rabh.). Type on Osyridicarpos natalensis. South Africa, Natal, Tabamhlope, leg. Wood No. 527.
Syn. Puccinia osyridicarpi (Massee) Grove in Wakefield & Grove 1916: 76.
Pycnia not seen.
Aecia aecidioid, amphigenous on leaves and caulicolous, scattered in + concentric groups up to 5 mm diameter, causing slight hypertrophy, without spots surrounding aecia, aecial cups small cylindrical, circa 250 um diameter, up to 500 um long, spore mass pale yellowish to white, surrounded by the irregularly and deeply frayed, orange- yellow to whitish peridium, cells of the peridium firmly connected, outer wall finely striate, circa 12-17 um wide, inner wall delicately and closely verrucous, circa 4—6 um wide. Aeciospores angular globoid, subgloboid to ovoid, 23—30 x 22—27 um, spore wall about 1—1.5 um thick, inconspicuously finely verrucous, hyaline, germ pores invisible.
Uredinia amphigenous on leaves and on stems, (partly) developing from the uredinia, subepidermal, roundish, ellipsoid or irregularly in outline, up to | mm wide, separate or scattered, without spots surrounding uredinia, cinnamon-brown, early exposed, pulver- ulent, surrounded by the torn epidermis. Urediniospores subgloboid, ovoid or ellipsoid, (26—) 29-37 x 22—30 um, borne singly on pedicels, spore wall uniformly about 2—3 um thick, yellowish-brown, noticeable verrucous, germ pores (3 to) 4 to 5, mostly + equa- torial, occasionally scattered, with hyaline papillae that are weakly developed.
Telia amphigenous on leaves and caulicolous, subepidermal, roundish, ellipsoid or ir- regularly in outline, up to 5 mm diameter, separate or scattered, blackish-brown to black, early exposed, compact to cushion-like, surrounded by the torn epidermis, with- out spots surrounding telia. Teliospores in general bicellular, obovoid, ellipsoid to ob- long, slightly constricted at the septum, 38-48 x 21—31 um, rounded or subacuminate at the apex, rounded or attenuate at the base, spore wall about 2-4 um thick at the sides,
26
about 6—9 um thick at the apex, brown, smooth, germ pore of the upper cell apical, germ pore of the lower cell just below the septum, without papillae, pedicel persistent, up to 100 um long, thick-walled, mostly not collapsing, yellowish to hyaline, sometimes ob- liquely inserted. Mesospores occasionally existing.
Fig. 17: Puccinia cf. stonemaniae on Thesium cf. strictum. Peridium cells and aeciospores (RSA 216), uredinio- and teliospores (RSA 3). Scale bar = 10 um.
Specimens examined:
- On Thesium cf. strictum Bergius. South Africa, Western Cape Province, Cape Peninsula National Park, BIO- TA-observatory at Olifantsbos, S 34° 15’ 45.3’’, E 18° 23’ 36.6’, 17.X1.2001, leg. M. Mennicken No. RSA 3, IJ Il] (PREM, M).
- On Thesium cf. strictum. South Africa, Western Cape Province, Cape Peninsula National Park, BIOTA-obser- vatory at Olifantsbos, S 34° 15’ 41.1’, E 18° 23’ 36.8’, 23.1X.2002, leg. M. Mennicken No. RSA 216, I (PREM, M).
- On Thesium viridifolium Levyns. South Africa, Western Cape Province, Cape Peninsula National Park, BIO- TA-observatory at Olifantsbos, S 34° 15’ 55.1’, E 18° 23’ 52.5’’, 23.1X.2002, leg. M. Mennicken No. RSA 217, II III (PREM, M).
Our collections do not agree very well with the description of Puccinia stonemaniae given in the diagnosis (Sydow & Sydow 1912) and in Doidge (1927). Therefore we use the epithet with the prefix cf. Unfortunately the type collection of Puccinia stonemaniae in Berlin got mouldy. Therefore, it could not be examined. Thesium viridifolium seems to be a new host plant.
Acknowledgements The authors thank Pedro W. Crous and Halvor B. Gjzrum for peer review of the manu- script, the curators of B, PREM, and WIND for the loan of specimens, Matthias Lutz,
2)
Wolfgang Maier, and Michael Wei8 for critical comments on the manuscript, Michael WeiB for help with the Latin diagnosis, and Sheila and Brian Longman for comments on the English text. The study was made possible by the Federal Ministry of Education and Research in Germany through the sourcing of the BIOTA Southern Africa project. The Northern Cape Department of Nature and Environmental Conservation, the Western Cape Nature Conservation Board, and the Conservation Services of the South African National Parks are gratefully acknowledged for issuing research permits. The authors thank all land owners and decision makers for the permission to enter the BIO- TA-observatories and to collect rust-infected plants. The botanical BIOTA working group of Norbert Jiirgens from the University of Hamburg is gratefully acknowledged for help with the determination of the host plants.
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MYCOTAXON
Volume 90, pp. 29-34 July-August 2004
New species of Asterina from Guangdong, China
BIN Sono*, Tar-Hut Li, YA-HENG SHEN
binsong @ gdas.ac.cn Guangdong Provincial Key Laboratory of Microbial Culture Collection and Application; Guangdong Institute of Microbiology Xianlie Road Central, Guangzhou 510 070,China
Abstract—Asterina daphniphyllicola parasitic on Daphniphyllum macropodum, Asterina dinghuensis parasitic on Cleistocalyx operculatus, Asterina euryae parasitic on Eurya acuminata, and Asterina myrsinacearum parasitic on Myrsinaceae are described and illustrated as new species.
Key words—Asterinaceae, Daphniphyllaceae, Myrtaceae, Theaceae
Introduction
Guangdong Province of China is Mainland China’s most southern region. It is close to Hainan Islands, and together they constitute the main tropical and subtropical region of the country. It is an area of abundant and diverse vegetation, characterized by warm, humid weather and frequent rainfall. It has one of the highest levels of fungal diversity of any area in China (Song et al. 2002, 2003). Recently, the first author had the opportunity to restudy the specimens of the asterinaceous fungi deposited in the Herbarium of Guangdong Institute of Microbiology (HMIGD), Guangzhou (China). The study shows that four taxa of the genus Asterina occurring in Guangdong are. They are apparently undescribed and are described and illustrated as new species as follows here.
Asterina daphniphyllicola B. Song, sp. nov. Fig. 1
Similis Asterina daphniphylli sed differt appressoriis et ascosporis brevioribus, appressoriis 10% oppostiis.
Etymology: daphniphyllicola, in reference to the host, Daphniphyllum macropodum.
Colonies amphigenous, mostly epiphyllous, black, thin, arachnoid to nearly velvety, scattered, up to 5 mm in diameter, sometime confluent. Hyphae brown, sinuous to nearly straight or mostly flexuous, irregularly branching acutely or widely, loosely or densely reticulate, cells mostly 18-30 x 4-5 wm. Appressoria unicellular, alternate to 10% opposite, spreading, straight or bent, ovate, obtuse or narrow at apex, entire to angular, sometime lobate, 7-10 x 5-7 wm. Ascomata scattered to nearly aggregate, black, orbicular to hemispherical, up to 230 wm in diameter, stellately dehiscent at the center, irregularly crenate to short tasseled at periphery, surface cells 2-3 wm wide. Ascospores
30
cylindrical or fusoid, narrow or obtuse at ends, brown, |-septate, constricted at septum, smooth, 25-33 x 8.5-10 ym.
Holotype: On leaves of Daphniphyllum macropodum Miq.(Daphniphyllaceae). Nankung Shan, Guangdong Provinice, China. September 17,1992, Y. S. Ouyang and B. Song, HMIGD 32067.
The new species is closely related to Asterina daphniphylli W. Yamam. The main distinguishing characters of A. daphniphylli are that the appressoria are longer (9-18 pm), alternate and not opposite, and the ascospores are broader (12-16 ~m) (Yamamoto 1956).
Asterina dinghuensis, sp. nov. Fig. 2
Similis Asterina assimilis sed differt appressoriis et ascosporis brevioribus; et a A. claviflori differt ascosporis grandioribus (20-28 x 10.5-12.5 jum).
Etymology: dinghuensis, in reference to the type locality, Dinghu Shan Biosphere Reserve.
Colonies amphigenous, thin, black, arachnoid or nearly velvety, up to 5 mm in diameter, sometimes confluent. Hyphae brown, sinuous or nearly straight, opposite or irregular branching acutely or obtusely, loosely to closely reticulate, cells mostly 20-45 x 3.8-5 pm. Appressoria unicellular, in alternate or unilateral arrangement, spreading, straight or slightly bent, cylindrical, narrow at apex, entire or sometimes angular, 10-16 x 4- 5.5 um. Ascomata scattered to nearly aggregate, black, orbicular or hemispherical, up to 260 wm in diameter, no apical pore or stellately dehiscent at center, irregularly crenate to shortly tasseled at periphery, surface cells 2-3.5 ~m wide. Ascospores nearly ellipsoid to oblong, brown, 1-septate, obtuse, constricted at septum, smooth, 20-28 x 10.5-12.5 wm.
Holotype: On leaves of Cleistocalyx operculatus (Roxb.) Merr. et Perry (Myrtaceae). Dinghu Shan Biosphere Reserve, Guangdong Province, China. December 14, 1978, G.Z. Jiang, HMIGD 30008.
The appressoria of this species are unicellular, alternate and not opposite, nearly cylindrical. This species is similar to Asterina assimilis Syd., Asterina clavifiori A. K. Kar et Maity, Asterina eugeniae H. S. Yates, Asterina natalitia Doidge, and Asterina rickii Theiss. (Theissen 1913; Yates 1917; Sydow 1922; Doidge 1942; Kar & Maity 1970). According to their descriptions, the main distinguishing characters for A. assimilis are that the ascospores and the appressoria are broader (24-26 x 13-15 wm for the ascospores, 10-17 x 7-9 wm for the appressoria), and for A. claviflori that the ascospores are smaller (9.9-13.2 x 6.6-9.9 wm). The main distinguishing characters for A. rickii are that the ascospores are narrower (18-22 x 8-9 wm), and for A. eugeniae that the ascospores are smaller (20 x 5 wm) and the appressoria are irregular. In addition, the new species differs from A. natalitia, which has larger ascospores (30-34 x 12.5-15 pm). Therefore, they can clearly be distinguished from the new species.
The new species is found mixed with Meliola leptospermi Hansf. (Hu et al. 1996).
6
Fig.1 Asterina daphniphyllicola (a. hyphae with appressoria; b. ascoma; c. ascospores)
Fig.2 Asterina dinghuensis (a. hyphae with appressoria; b. ascoma; c. ascospores) (Bars = 25 ym).
52
Asterina euryae B. Song, sp. nov. Fig. 3
Similis Asterina theae sed differt ascosporis et appressoriis minoribus. Etymology: euryae, in reference to the host, Eurya acuminata.
Colonies epiphyllous, thin, black, arachnoid to nearly velvety, up to 3 mm in diameter, sometimes confluent. Hyphae brown, nearly straight or curved, branching acutely or obtusely, loosely to closely reticulate, cells mostly 10-25 x 7-8 wm. Appressoria unicellular, in alternate or unilateral arrangement, spreading, straight or bent, globose, mostly angular or sublobate, 12-13 x 11-12.5 wm. Ascomata scattered to nearly aggregate, black, orbicular to hemispherical, up to 240 ym in diameter, stellately dehiscent at the centre, crenate to shortly tasseled at periphery, surface cells 2.5-3.5 pm wide. Ascospores oblong, brown, 1-septate, obtuse, constricted at septum, surface smooth, 30-33 x 11.5-12.5 jum.
Holotype: On leaves of Eurya acuminata DC (Theaceae). Ruyang County, Guangdong Province, China. December 5, 1995, B. Song, HMIGD 34169.
The new species is closely related to Asterina theae W. Yamam. The main distinguishing characters of A. theae are that the ascospores and the appressoria are larger (35-44 x 14- 17 ym for the ascospores, 11-18 x 12-17 wm for the appressoria) (Yamamoto 1957).
Asterina myrsinacearum B. Song, sp. nov Fig. 4
Similis Asterina aphanes sed differt appressoriis grandioribus (10-19 x 8-12 um) et ascosporis Jusoideiis vel subellipsoideiis; et a A. embeliae differt ascosporis grandioribus (17-25 x 10- 13 yum).
Etymology: myrsinacearum, in reference to the host, Myrsinaceae indet.
Colonies epiphyllous, black, thin to nearly dense, arachnoid to nearly velvety, up to 8 mm in diameter, sometimes confluent. Hyphae brown, nearly straight to slightly sinuous, opposite or irregular branching acutely or obtusely, loosely or closely reticulate, cells mostly 17-30 x 3.5-5 wm. Appressoria unicellular, in alternate or unilateral arrangement, less than | % opposite, spreading, mostly bent, nearly globose to oblong, 1-3-sublobate or sometimes angular, obtuse at apex, 10-19 xX 8-12 wm. Ascomata nearly aggregate to scattered, black, orbicular or hemispherical, up to 130 wm in diameter, stellately dehiscent at the centre, crenate to shortly tasseled at periphery, surface cells 1.8-2.8 wm wide. Ascospores fusoid to nearly ellipsoid, brown, 1-septate, obtuse or narrowed at the ends, constricted at septum, surface smooth, 17-25 x 10-13 jum.
Holotype: On leaves of Myrsinaceae indet. Yangchun County, Guangdong Province, China. September 1986, H. Hu, HMIGD 34437.
The new species is closely related to Asterina aphanes Petr., Asterina ardisiae Hanstf., and Asterina embeliae Hansf. (Sydow & Petrak 1929; Hansford 1943, 1954). The main distinguishing characters of A. aphanes are that the appressoria are smaller (6-8 x 7-8 pm), and the ascospores are oblong-clavate. The main distinguishing characters of A. ardisiae are that the appressoria are cylindrical (8-14 x 6-8 jm), attenuate at apex, and the ascospores are oblong. The new species differs from A. embeliae in the latter having larger ascospores (26-28 X 13-15 wm).
a euryae (a. hyphae with appressoria; b. ascoma; c. ascospores) myrsinacearum (a. hyphae with appressoria; b. ascoma; c. ascospores) (Bars = 25 ym).
Fig.3 Asterin
Fig.4 Asterina
34 Acknowledgements
The authors wish to thank Prof. Dr. R. D. Goos of Department of Botany, University of Rhode Island (U.S.A.) and Prof. Dr. Richard P. Korf for serving as pre-submission reviewers and for their valuable comments and suggestions. They also acknowledge Prof. S Q Chen of South China Institute of Botany, Academia Sinica, Guangzhou, for identifying the host plants, and Ms. A.L. Zhang for technical assistance. This project was supported by the National Science Foundation of China (no. 30370012), the Natural Science Foundation of Guangdong (020470), the Natural Science Fund for Distinguished Young Scholars from Guangdong Academy of Sciences, and the Foundation of Guangdong Institute of Microbiology.
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Hansford CG. 1944. Contributions towards the fungas flora of Uganda. « 6 ». New records. Proc. Linn. Soc. Lond. 156:102-124.
Hansford CG. 1954. Some Microthyriales and other fungi from Indonesia. Reinwardtia 3:113-
144.
Hu YX, Ouyang YS, Song B, Jiang GZ. 1996. Meliolales « 1 ». Flora Fung. Sin. 4:1-270 (in Chinese)
Kar AK, Maity MK. 1970. New Asterina spp. from West Bengal. Trans. Br. Mycol. Soc. 54:435- 444.
Song B, Li TH, Shen YH. 2002. Ecological and Floristic analyses of Meliolales (Fungi) in China. Journ. Trop. Subtrop. Bot. 10(2):118-127.
Song B, Li TH, Hosagoudar VB. 2003. Four new Asterina species from Yunnan, China. Fungal Diversity 14:157-164.
Sydow H. 1922. The Amboina Fungi Collected by C. B. Robinson. Philipp. Journ. Sci. C. Bot. 21:131-146.
Sydow H, Petrak F. 1929. Fungi costaricenses a cl. prof. Alberto M. Brenes collecti series prima. Ann. Mycol. 27:1-86.
Theissen F. 1913. Die Gattung Asterina in Systematischer Darstellung Abh. der K.K. Zoologisch- Bot. Gesell. Wien. 7:1-138,pl.1-8.
Yamamoto W. 1956. The Formosan Species of the Microthyriaceae « | ». Sci. Rep. Hyogo Univ. Agr. Ser. Agr. Biol. 2:33-36.
Yamamoto W. 1957. The Formosan Species of the Microthyriaceae « 2 ». Sci. Rep. Hyogo Univ. Agr. Ser. Agr. Biol. 3:23-32
Yates HS. 1917. Some recently collected Philippine Fungi « 1 ». Philipp. Journ. Sci. C. Bot. 12:361-380.
MYCOTAXON
Volume 90(1), pp. 35-42 July-September 2004
Preliminary survey of the Helvellaceae from Xinjiang, China
WEn- YING ZHUANG!
zhuangwy @sun.im.ac.cn Key Laboratory of Systematic Mycology & Lichenology Institute of Microbiology, Chinese Academy of Sciences Beijing 100080, China
Abstract -- Collections of the Helvellaceae from Xinjiang, China were examined. 17 taxa of the genera Gyromitra, Helvella, Hydnotrya, and Wynnella are recognized. Among them, H. cupuliformis var. crassa and H. jimsarica are new taxa. Helvella cupuliformis var. cupuliformis is new to China. Distinctions between the new taxa and their closely related species are discussed.
Key words -- taxonomy, Gyromitra, Helvella, Hydnotrya, Wynnella
Introduction
Xinjiang is in the northwest of China. It occupies more than 1/7 of the China mainland territory (Fig. 1). The area is covered with deserts, grasslands, conifer forests, broadleaf trees, and shrubs. The Tianshan Mountains run east-west in the area and separate it into two parts, North Xinjiang and South Xinjiang. Mt. Altay is located in the very north and Mt. Kunlun is at the southwest border. The Junggar Basin lies between Mt. Altay and the Tianshan Mountains. Tarim Basin lies between the Tianshan Mountains and Mt. Kunlun. The huge Taklimakan Desert is in the center of Tarim Basin. The mean annual precipitation varies from 50 mm to 600 mm. The mean annual temperature of flatlands is 6—8C in the north, 10-11C in the south; while that in the mountainous area is around 2.5-5C. Glaciers and snow cover the high peaks all the year round. Forests in Xinjiang mainly consist of Picea, Larix, Abies, Pinus, Juniperus, Betula, Populus, Hippophae, Salix, Haloxylon, Tamarix, Calligonum, Nitraria, etc. (Lu & Yan 1989).
A few helvellaceous cup-fungi were previously recorded from Xinjiang (Zhao & Mao 1986; Cao 1988; Cao et al. 1990a, b; Zhang 1990; Mao 1998).
' Supported by the National Natural Science Foundation of China (No. 30230020).
36
Recent collections of the Helvellaceae from Xinjiang were studied as were specimens of the group on deposit in the Mycology Herbarium, Chinese Academy of Sciences (HMAS). Thirteen taxa of Helvella, 2 of Gyromitra, | of Hydnotrya, and | of Wynnella are recognized. Xinjiang is rich in Helvella taxa and has high species diversity accounting for 40% of the total known species in China (Zhuang 1998). Helvella costifera, H. crispa, H. cupuliformis var. cupuliformis, H. ephippium, and H. macropus are the common species there. Helvella cupuliformis var. cupuliformis is a new record for China. Helvella cupuliformis var. crassa and H. jimsarica are described as new taxa. Gyromitra is also frequently encountered there. Collections of the genus from the eastern Xinjiang are mostly G. infula, while those from the west belong to G. xinjiangensis. Hydnotrya seems not to be widely dispersed. Wynnella is possibly common in Xinjiang.
Taxonomy
Gyromitra infula (Schaeff.) Quél., Ench. Fung. p. 272, 1886.
Specimens examined: CHINA. Xinjiang, Tuomuerfeng, VIII 1978, on rotten wood, S.X. Sun, H.A. Wen & X.L. Mao 597, HMAS 39333; Tianchi, alt. 1900 m, 31 VII 2003, on rotten wood, mycological team WYZ 4603, 4604, HMAS 86061, 86060; Jimsar, alt. 1700 m, 1 VUI 2003, on rotten wood, W.Y. Zhuang & Y. Nong 4659, HMAS 86057; Burgin, Hemuxiang, alt. 1100 m, 5 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4692, 4693, HMAS 86059, 86058; Burgin, Hemuxiang, alt. 1100 m, 6 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4731, 4732, HMAS 86056, 86055; Altayshan, alt. 1250 m, 9 VII 2003, on the ground, W.Y. Zhuang & Y. Nong 4820, HMAS 86054; Altayshan, alt. 1250 m, 9 VII 2003, on the ground, W.Y. Zhuang & Y. Nong 4829, HMAS 86049.
Gyromitra xinjiangensis J.Z. Cao, L. Fan & B. Liu, Acta Mycol. Sin. 9: 105, 1990.
Specimens examined: CHINA. Xinjiang, Hejing, | VII 1958, on rotten wood, L.W. Xu 132, HMAS 27863; Tuomuerfeng, 11 VII 1978, on the ground, S.X. Sun, H.A. Wen & X.L. Mao 760, HMAS 39091; Tuomuerfeng, 11 VII 1978, on rotten wood, S.X. Sun, H.A. Wen & X.L. Mao 598, HMAS 39334; Xinjiang, VIII 1994, on rotten wood, J.Y. Wang 310, HMAS 71869; Yining, Qapgal, alt. 2000 m, 13 VIII 2003, on rotten wood, W.Y. Zhuang & Y. Nong 4894, 4905, HMAS 86048, 86047; Xinyuan, Nalati, alt. 2200 m, 15 VUI 2003, on the ground, W.Y. Zhuang & Y. Nong 4964, 4968, HMAS 86046, 86053; Xinyuan, Gongnaisi, alt. 2170 m, 16 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4981, HMAS 86052.
Notes: This species is very similar to Gyromitra infula in gross morphology but different from the latter in the ascospore surface morphology, spore guttulation, and fruitbodies with a relatively short stipes (Cao et al. 1990a).
bi
Fig. | Map of China showing location of Xinjiang (left), map of Xinjiang showing collecting sites of the helvellaceous fungi (right).
Figs. 2-7 Morphology of Helvella spp. Figs. 2-4 Helvella cupuliformis var. crassa (paratype): 2. Dried fruitbodies, x0.75; 3. Ectal excipular structure, x165; 4. Ascospores, x330. Figs. 5-7 Helvella jimsarica (holotype): 5. Dried fruitbodies, x0.4; 6. Ectal excipular structure, x165; 7. Ascospores, x330.
38
Helvella acetabulum (L.) Quél., Ench. Fung. p. 275, 1886.
Specimens examined: CHINA. Xinjiang, Tuomuerfeng, 27 VI 1977, on the ground, H.A. Wen & X.L. Mao 53, HMAS 38329; Tuomuerfeng, 29 VI 1977, on the ground, H.A. Wen & X.L. Mao 56, HMAS 38129; Tianchi, alt. 1900 m, 31 VII 2003, on the ground, mycological team WYZ 4614, HMAS 86051; Jimsar, alt. 1700 m, 1 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4649,
HMAS 86050.
Helvella atra Holmskj. : Fr., Beata Ruris Otia Fungis Danicis Impensa 2: 47, bo
Specimens examined: CHINA. Xinjiang, Burgin, Hemuxiang, alt. 1100 m, 5 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4695, 4727, HMAS 86045, 86044.
Helvella costifera Nannf., in Lundell & Nannfeldt, Fungi Exsic. Suec. Praes. Upsal. Fasc. 41-42: 37, no. 2061, 1953.
Specimens examined: CHINA. Xinjiang, Tianchi, alt. 1900 m, 31 VII 2003, on the ground, mycological team WYZ 4600, 4616, HMAS 86043, 86042; Jimsar, alt. 1700 m, 1 VIL 2003, on the ground, W.Y. Zhuang & Y. Nong 4638, 4661, HMAS 86041, 86040; Jimsar, alt. 1700 m, 2 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4662, 4667, 4669, 4679, HMAS 83506, 83507, 83508, 83509; Hemuxiang, alt. 1100 m, 5 VIII 2003, on the ground, W. Y. Zhuang & Y. Nong 4684, HMAS 83510; Guozigou, alt. 1800 m, 11 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4864, 4865, HMAS 83511, 83512.
Helvella crispa (Scop.) Fr., Syst. Mycol. 2: 14, 1822.
Specimens examined: CHINA. Xinjiang, Jimsar, alt. 1700 m, 2 VIII 2003, on the ground, W.-Y. Zhuang & Y. Nong 4666, HMAS 83513; Burgin, Hemuxiang, alt. 1100 m, 5 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4697, 4696, HMAS 83514, 83550; Altayshan, alt. 1250 m, 9 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4821, 4822, HMAS 83515, 83516; Guozigou, alt. 1800 m, 11 VII 2003, on the ground, W.Y. Zhuang & Y. Nong 4867, HMAS 83517; Yining, Qapgal, alt. 2000 m, 13 VII 2003, on the ground, W.Y. Zhuang & Y. Nong 4898, HMAS 83518.
Helvella cupuliformis Dissing & Nannf., Sv. Bot. Tidskr. 60: 326, 1966. var. cupuliformis
Specimens examined: CHINA. Xinjiang, 15 VII 1978, on the ground, S.X. Sun, H.A. Wen & X.L. Mao 421, HMAS 38330; 23 VII 1977, on the ground, H.A. Wen & X.L. Mao 137, HMAS 38331, Tianchi, alt. 1900 m, 31 VII 2003, on the ground, mycological team WYZ 4605, 4606, 4618,
HMAS 83519, 83520, 83521; Jimsar, alt. 1700 m, 1 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4646, HMAS 83522.
Helvella cupuliformis var. crassa W.Y. Zhuang, var. nov. Figs. 2-4
Ab Helvella cupuliformis var. cupuliformis ascosporis crassis, 18-21 x 12.5-15 um differt.
Pileus discoid, 1.5-5.5 cm in diam., hymenium surface grayish brown to grayish, receptacle surface pubescent, pale gray, stipe terete, short, nearly smooth, beige; ectal excipulum of textura angularis, 115-130 um thick excluding pustules, outermost cells thin-walled, hyaline, club-shaped, 18-31
39
x 7.5-20 um; medullary excipulum of textura intricata, hyphae hyaline, 2.5-5 um wide; subhymenium not clearly defined; hymenium 230-245 um thick; asci subcylindrical, 8-spored, J— in Melzer’s reagent, 18-20 um wide; ascospores broadly ellipsoid, uniguttulate, 18-21 x 12.5-15 um; paraphyses somewhat enlarged at apex, 4.5—6 um wide at apex.
Holotype: CHINA. Xinjiang, Tianchi, alt. 1900 m, 31 VII 2003, on the ground, mycological team
WYZ 4615, HMAS 83523. Paratype: Xinjiang, Jimsar, alt. 1700 m, 1 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4648, HMAS 83524.
Notes: The new variety is very similar to Helvella cupuliformis var. cupuliformis in gross morphology and anatomic structure but differs significantly in the large fruitbodies and much wider ascospores (18-21 x 12.5-15 um vs. 18-20 x 11.5-13 um) (Dissing 1966).
Helvella elastica Bull., Herb. France Pl. 242, Figs. A, B, D-E, 1785. = Helvella xinjiangensis J.Z. Cao, L. Fan & B. Liu, Acta Mycol. Sin. 9: 186, 1990.
Specimens examined: CHINA. Xinjiang, Tuomuerfeng, 27 VII 1978, X.L. Mao et al. 447, HMAS 38353 (holotype of H. xinjiangensis); Jimsar, alt. 1700 m, 1 VII 2003, on the ground, W. Y. Zhuang & Y. Nong 4641, HMAS 83525; Mt. Altay, alt. 1250 m, 9 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4835, HMAS 83526.
Notes: Significant distinctions were not found between the holotype of Helvella xinjiangensis and H. elastica (Dissing, 1966, Cao et al., 1990b).
Helvella ephippium Lév., Ann. Sci. Nat. Ser. 2, 16: 240, 1841.
Specimens examined: CHINA. Xinjiang, Hejing, 2 VIII 1958, on the ground in woods, L.W. Xu 153, HMAS 27866; Tuomuerfeng, 15 VII 1978, on the ground in woods, X.L. Mao et al. 423, HMAS 38354; Tianchi, alt. 1900 m, 31 VIL 2003, on soil, fungus collecting team, WYZ 4601, HMAS 83527; Jimsar, alt. 1700 m, 1 VII 2003, on the ground, W.Y. Zhuang & Y. Nong 4639, HMAS 83528; Guozigou, alt. 1800 m, 11 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4870, HMAS 83529; Yining, Qapqal, alt. 2000 m, 13 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong
4923, HMAS 83530.
Helvella jimsarica W.Y. Zhuang, sp. nov. Figs. 5-7
Ab Helvella elastica apotheciis lignicola, grandiebus, superficiebus receptaculo pustulatis, pubescens vel villosis, subhymeniis tennuibus differt.
Pileus discoid to somewhat saddle-shaped, up to 5.5 cm in diam. and 8 cm high when fresh, hymenium surface beige to brown when fresh and dark brown when dry, receptacle surface pubescent to villose, often with minute pits observed when dry, light grayish brown to grayish brown when fresh and grayish cinnamon when dry, stipe terete, with a pubescent to villose surface, beige when fresh and straw-colored to grayish beige when dry; ectal excipulum of textura angularis, 75-100 um thick excluding the pustules, pustules 25-80 um high,
40
outermost cells thin-walled, club-shaped, (12—)15-38 x 7.5-21 um, arranged in chains to give rise to the pustules; medullary excipulum of textura intricata, 330-600 um thick or thicker, hyphae hyaline, 3-7.5 um wide; subhymenium ca 50 um thick; hymenium 250-280 um thick; asci subcylindrical, 8-spored, J— in Melzer’s reagent, 13-16.5 um wide; ascospores ellipsoid to broadly ellipsoid, some with warts on surface, uniguttulate, uniseriate, 16.5-21 x 10-13 um; paraphyses enlarged to 6—7 um wide at apex.
Holotype: CHINA. Xinjiang, Jimsar, alt. 1700 m, 1 VIII 2003, on rotten wood, W.Y. Zhuang & Y.
Nong 4660, HMAS 83531. Paratype: CHINA. Xinjiang, Jimsar, alt. 1700 m, 1 VIII 2003, on plant debris, W.Y. Zhuang & Y. Nong 4640, HMAS 83532.
Notes: The hymenium surface color of Helvella jimsarica when fresh resembles that of Helvella elastica, but the surfaces of receptacle and of stipe are entirely different from those in H. elastica. The new species is characterized by the very large fruitbodies up to 5.5 cm in diameter and 8 cm high, pubescent to villose receptacle surface light grayish brown to grayish brown, with minute pits on surface when dry, as well as its occurrence on rotten wood and plant debris.
Helvella lacunosa Afz. : Fr., Kgl. Vet.-Akad. Nya Hand. 4, p. 303, 1783. = Helvella sulcata Afz. Kgl. Vet.-Akad. Nya Hand. 4, p. 305, 1783.
Specimens examined: CHINA. Xinjiang, Tuomuerfeng, on the ground in woods, 24 VII 1977, X.L. Mao 158, HMAS 38355; Jimsar, alt. 1700 m, 1 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4628, 4642, HMAS 83533, 83537; Jimsar, alt. 1700 m, 2 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4663, 4678, HMAS 83534, 83535; Burgin, Hemuxiang, alt. 1100 m, 5 VIII
2003, on the ground, W.Y. Zhuang & Y. Nong 4694, HMAS 83536.
Notes: HMAS 83537 is somewhat different from other collections. The gross morphology of the fungus is very similar to that of typical Helvella lacunosa. It is distinguished from Nordic (a), British (b), and other Chinese (c) collections in wider asci [15-20 um vs. 13-16 um (a), ca 18 um (b), and 15-18 um (c) wide], larger ascospores [18-—21.5 x 11.5-14.5 um vs. 15-17.1 x 9.5-12 um (a), 17-20 x 11-13 um (b), and 15-20 x 11.5-13 um (c)], and lack of a well- developed subhymenium (Dissing 1966, Dennis 1978). Since spore sizes of specimens from the different geographical areas do not differ clearly, the above distinctions are treated as variations currently unworthy of taxonomic rank. The morphology of HMAS 83537 is described as follows.
Pileus saddle-shaped, 2-5 um wide and 4.5-—9 um high, hymenium surface grayish brown, receptacle surface paler and smooth to slightly pubescent, stipe robust, with longitudinal furrows, pale grayish to nearly black when dry; ectal excipulum of textura angularis, 130-155 um thick, cells thin-walled, forming a palisade outer layer, 20-38 x 13-20 um; medullary excipulum
4]
of textura intricata, 280-380 um thick or thicker, hyphae hyaline, 2-5 um wide; subhymenium not clearly recognized; hymenium 305-325 um thick; asci subcylindrical, (4—)8-spored, J- in Melzer’s reagent, 15-20 um wide; ascospores broadly ellipsoid, uniguttulate, uniseriate, 18-—21.5 x 11.5-14.5 um; paraphyses enlarged at apex, 5—7.5(—8) um wide near apex and 3-3.5(-4) um wide below.
Helvella macropus (Pers. : Fr.) P. Karst., Not. Sallsk. F Fl. Fenn. Forh. 11: 224, 1870.
Specimens examined: CHINA. Xinjiang, Tianchi, alt. 1900 m, 31 VII 2003, on the ground, mycological ttam WYZ 4617, HMAS 83538; Jimsar, 1700 m, 1 VII 2003, on duff, W.Y. Zhuang & Y. Nong 4626, HMAS 83539; Jimsar, 1700 m, 2 VIII 2003, on duff, W.Y. Zhuang & Y. Nong 4674, HMAS 83540; Burgin, Hemuxiang, alt. 1100 m, 5 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4698, HMAS 83541; Guozigou, alt. 1800 m, 11 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4851, 4868, 4869, HMAS 83542, 83543, 83544.
Helvella phlebophora Pat. & Doass., in Pat., Tab. Anal. Fung. 5: 208, 1886.
Specimen examined: CHINA. Xinjiang, Burgin, Hemuxiang, alt. 1100 m, 6 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4774 , HMAS 83545.
Helvella spadicea Schaeff., Fung. Icon. 3, Tab. 283, 1772; l.c. 4: 112, 1774. = Helvella leucopus Pers., Mycol. Eur. 1: 213, 1822.
Specimens examined: Bachu, VI 1985, on the ground, C. Gao 1661, HMAS 50932; Bachu, V 1986, Anon. 1904, HMAS 53650; V 1987, on the ground I broadleaf forest, Z.Y. Zhao, HMAS 57640; Kashi, on the ground in broadleaf forest, Q. Ji, HMAS 72884.
Notes: The fungus was previously recorded in the Chinese literature as Helvella leucopus (Mao 1998).
Hydnotrya cerebriformis Harkn., Proc. Calif. Acad. Sci. 3, 1: 266, 1899. Specimen examined: CHINA. Xinjiang, Xinyuan, Nalati, alt. 2200 m, 15 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4954, HMAS 83546.
Notes: According to Zhang (1990), the fungus was also collected in Qitai County in the east of Xinjiang.
Wynnella auricula (Schaeff.) Boud., Hist. Class. Discom. Eu. p. 51, 1907. = Elvela auricula Schaeff., Fung. Bavar. Ind. p. 103, 1774.
= Wynnella atrofusca (Beck) Svrcek, Ceska Mykol. 17: 45, 1963. = Otidea atrofusca Beck, flora Herstein S. A. p. 131, 1885.
= Wynnella silvicola (Beck in Sacc.) Nannf., Ann. Bot. Fenn. 3: 309, 1966. = Otidea silvicola Beck in Sacc., Syll. Fung. 8: 97, 1889.
Specimens examined: CHINA. Xinjiang, Jimsar, alt. 1700 m, 1 VIII 2003, on rotten wood, A. Abbas, W.Y. Zhuang & Y. Nong 4668, HMAS 86062; Yining, Qapqal, alt 2000, 13 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4924, HMAS 83547; Guozigou, alt. 1800 m, 11 VIII 2003, on the ground, W.Y. Zhuang & Y. Nong 4866, HMAS 83548.
42
Notes: The fungus has been found in North America, Europe, and China (Seaver 1928, Dissing 1966, Eckblad 1968, Hansen & Knudsen 2000). It obviously has north temperate distribution. It was previously collected from Mt. Guancen in Shanxi Province, Urmuqi in Xinjiang, and Mt. Yulong in Yunnan (Cao 1988).
Acknowledgements
The author would like to thank Prof. R. P. Korf and Prof. Z. Z. Li for serving as the pre-submission reviewers, Mr. Y. Nong and other members in the mycological team for collecting jointly the specimens used in this study, Mr. S. X. Sun for technical assistance, Ms. X. Song for making the sections, and Mr. X. P. Zhao for helping with the photographs.
Literature Cited
Cao JZ. 1988. Study on the Helvellaceae from China. MS Thesis. Shanxi University, Taiyuan.
Cao JZ, Fan L, Liu B. 1990a. Notes on the genus Gyromitra from China. Acta Mycol. Sin. 9: 100- 108. (in Chinese)
Cao JZ, Fan L, Liu B. 1990b. Some new species and new records of the genus Helvella from China. II. Acta Mycol. Sin. 9: 184-190. (an Chinese)
Dennis RWG. 1978. British Ascomycetes. Ed. 2. Cramer, Vaduz.
Dissing H. 1966. The genus Helvella in Europe with special emphasis on the species found in Norden. Dansk. Bot. Arkiv 25(1): 1-172.
Eckblad F-E. 1968. The genera of the operculate discomycetes — A re-evaluation of their taxonomy, phylogeny and nomenclature. Nytt Mag. Bot. 15: 1-191.
Hansen L, Knudsen H. 2000. Nordic Macromycetes. Vol. 1. Nordsvamp, Copenhagen.
Kirk PM, Cannon PF, David JC, Stalpers JA. 2001. Ainsworth & Bisby’s Dictionary of the Fungi.
CAB International, Surrey UK.
Lu P, Yan GX. (eds.) 1989. Forests of Xinjiang. Xinjiang People Press, Urmuqi. China Forestry Press, Beijing. (in Chinese)
Mao XL. 1998. Economic Fungi of China. Science Press, Beijing. (in Chinese)
Seaver FJ. 1928. The North American Cup-fungi (Operculates). Seaver. New York.
Zhang BC. 1990. Systematics of Hypogeous Pezizales and Taxonomy of the Chinese Genera and Species. Ph D Thesis, Institute of Microbiology, Chinese Academy of Sciences, Beijing. (in Chinese)
Zhao ZY, Mao XL. 1986. Jcones of Macrofungi from Xinjiang. Xinjiang Bayi Agricultural College, Urmudi. (in Chinese)
Zhuang WY. 1998. A list of discomycetes in China. Mycotaxon 67: 465-390.
MYCOTAXON
Volume 90(1), pp. 43-54 July-September 2004
Genetic variability in tropical and temperate populations of Trapeliopsis glaucolepidea: Evidence against long-range dispersal in a lichen with disjunct distribution
ZDENEK PALICE* & CHRISTIAN PRINTZEN**
* palice @ibot.cas.cz Institute of Botany, Academy of Sciences of the Czech Republic CZ-25243 Pruhonice, Czech Republic
** christian.printzen @senckenberg.de Forschungsinstitut Senckenberg, Abt. Botanik/Paldobotanik
Senckenberganlage 25, D-60325 Frankfurt am Main, Germany
Abstract—The squamulose lichen Trapeliopsis glaucolepidea displays a widely disjunct distribution in Europe, East Africa, tropical America, and New Guinea. Using ITS sequences from 38 European and 31 Latin American individuals, we compared the genetic variability within and among populations of this species. Specifically, we addressed the questions whether recurrent genetic exchange occurs between Latin America and Europe, whether there are signs of historical long-range dispersal, whether genetic variability is higher in tropical than in temperate populations, and whether the separation of T. glaucolepidea and T. percrenata is supported by genetic characters. Intercontinental gene flow seems to be absent and there are also no signs of historical long-range dispersal. Two deviating haplotypes from Europe might belong to an unrecognized species or indicate the presence of paralogous ITS sequences. But our dataset does not support separation of T. glaucolepidea and T. percrenata based on morphological characters. More thorough sampling efforts and more sequences are necessary to make more detailed inferences on the phylogeography of T: glaucolepidea.
Key words—genetic diversity, lichenized fungi, nuclear DNA, Ecuador, Czech Republic
Introduction
Phylogeography provides powerful tools for the study of historical biogeography, dispersal biology, and speciation in various organismal groups (e.g. Avise 2000, Templeton 1998, 2001). Since direct observations are impossible inferences about a species’ history can only be made based on genetic data. Population genetics of lichens is still largely unexplored and thus most questions concerning the dispersal, biogeography or speciation of lichens are still unanswered (Printzen & Ekman 2002). Especially the wide disjunctions of many lichen species as compared to other organisms, such as phanerogams are still unexplained. Fragmentation of formerly coherent distributional ranges and long-range dispersal were both invoked as explanations. Some of the first published studies on infraspecific genetic variability of lichens concentrated on biogeographical questions, but came to deviating conclusions. Printzen et al. (2003)
44
inferred that the North American-European disjunction of Cavernularia hultenii was due to the postglacial fragmentation of a circumarctic distributional area, while H6gberg et al. (2002) concluded that European populations of Letharia vulpina were the result of long range dispersal from North America. In this study we try to provide data on the genetic variability of another lichen species, this time with a peculiar tropical-European disjunction.
Trapeliopsis glaucolepidea (Nyl.) G.Schneider (Ascomycotina, Agyriales) is a widespread squamulose lichen. Outside Europe, it is reported from higher elevations (mostly above 3000 m) in the tropics: East Africa (Swinscow & Krog 1988), Central & Southern America (Sipman 1999) and Papua New Guinea (Aptroot & Sipman 1991). A few additional unpublished collections exist from montane areas in Asia and a middle-montane region in South America (Palice, unpubl.). No North American records are known so far. In Colombia (Cleef in sched., cf. Sipman et al. 2000) and Ecuador (Palice, pers. obs.) 7. glaucolepidea appears to be one of the most prominent cryptogamic components of the wet high-paramo — a mountain ecosystem characteristic for the Northern Andes. It grows mainly on naked peat, soil, dry grass tussocks, decaying rosettes of Asteraceae, Apiaceae and other kinds of plant debris. More rarely, it may be found on rocks or bark covered by mosses or decaying wood. In Europe, the species is best developed in the British Isles, where it preferably occupies open moorland areas at higher elevations. In continental Europe, 7. glaucolepidea prefers boreal coniferous forests, but also occurs in humid mixed/deciduous forests. Within one locality the species may grow on different substrates: soil, wood, shaded rock and plant debris. The most favoured substrate is relatively freshly exposed, naked peat, e.g. among roots of upended trees. It may also grow epiphytic at the bases of trees (Hafellner 2002).
T. glaucolepidea is a morphologically variable species. Andean and European populations differ in thallus size and in the production of apothecia. In general, the tropical specimens are larger (forms with small squamules are very rare) and more frequently fertile. Well developed tropical individuals can form large palmate squamules exceeding | cm in diameter. The best developed specimens from W- European moorlands are morphologically rather similar to average individuals from the tropics forming characteristic grey squamules c. 1-4 mm broad with bluish-grey labriform marginal soralia. Wood-inhabiting European specimens, on the other hand, may have squamules of the breadth only 0.3-1.0 mm. As they may produce soralia at a very early stage it can be difficult to distinguish squamules at all. In Central Europe, a continuum of intermediate forms between the extremes occurs. Thalli of smaller shade forms often lack the bluish tinge and the soralia are pale green to creamy white. Richly fertile specimens are only sparsely sorediate or the soredia are missing completely. The tropical material and the upland specimens from the British Isles have comparatively large apothecia, often above | mm in diameter while the specimens from forests produce the apothecia more rarely and they are usually smaller, rarely exceeding 1 mm in diameter. The above mentioned morphological differences apparently reflect environmental conditions, such as light, humidity, character of substrate (texture, decay rate, disturbance of surface) and perhaps also intra- and interspecific competition with other organisms.
Small specimens from decaying wood in West/Central European woodlands are usually
45
called Trapeliopsis percrenata. Based on statements on Scandinavian material (cf. Muhr 1986), observations from Bohemia and the respective herbarium material, Palice (1999) concluded that 7: percrenata is conspecific with T: glaucolepidea. In the South American paramo, sorediate specimens and richly fertile individuals without soralia occur. Their status is often difficult to assess. In addition to sparsely sorediate forms of T. glaucolepidea, non-sorediate specimens with potential affinities to the African T. haumanii or the Australasian 7. colensoi (indicated by ITS data not included in the current study) might be present.
This study attempts a preliminary answer on three main questions. (1) Is there evidence for recurrent or historical genetic exchange between Latin American and European populations of T. glaucolepidea? (2) Are there differences in the genetic diversity of tropical vs. temperate populations. (3) Is there evidence that morphologically different specimens (previously referred to 7. glaucolepidea and T. percrenata) really belong to two different species. Particularly with regard to this last question, but also in order to capture as much of the genetic diversity of 7. glaucolepidea as possible, our sampling tries to cover the complete morphological variability of European and Latin American T. glaucolepidea.
Materials and Methods
Sampling
We sampled 38 European and 31 Latin American individuals of 7: glaucolepidea (Append. 1) trying to cover the morphological variation of this species. For the comparison of genetic variability in tropical and temperate populations, we sampled several specimens from nearby populations in approximately equal areas of Ecuador (as a hypothetical diversity center) and the Czech Republic. In Ecuador, we chose four closely situated populations at Volcan Chiles, prov. Carchi (E1, n=17), a population in the NP Llanganates, prov. Napo/Tungurahua (E2, n=7), and one in the NP Podocarpus, Loja prov. (E3, n=4) for more intensive sampling. In the Czech Republic two populations in the NP Sumava, S-Bohemia (C1, n=12) and the NP Ceské Svycarsko, N-Bohemia (C3, n=7) were selected (Tab. 3, cf. Append. 1). The size of the populations varies somewhat within our dataset but was chosen so that the most distant collections from one population are not situated more than 50 km from each other. Only specimens that were clearly assignable to T. glaucolepidea were sampled, ITS sequences of non- sorediate specimens were only included when they belonged to one of the “sorediate” haplotypes.
DNA-sequencing
DNA was extracted from the vegetative lichen thalli using the NucleoSpin® Plant Kit (Macherey Nagel) according to the manufacturer’s protocol. In order to minimize contamination by other fungi — when feasible — younger thalli were used, and cortex and soralia were cut-off. PCR amplification of the internal transcribed spacer of the nuclear ribosomal DNA repeat (ITS) was performed in 50 wL volumes containing 3- 4 pL DNA, 5 vL 10 x reaction buffer, 2.5 mM MgCl, 1 mM dNTPs, 1 U Zag (MBI Fermentas), and 0.8 uM each of the primers ITS | F-5° (Gardes & Bruns 1993) and ITS 4-3° (White et al. 1990). Cycling conditions were as follows: initial denaturation at 94°C (4’), 6 cycles of a touch-down PCR with 94°C (1°), 62°C (-1°C per cycle) (1’), 72°C
46
(1°45’’), 34 cycles with 94°C (30), 56°C (30), 72°C (1’45”’’), and a final extension of 72°C (10’). PCR products were purified with the QIAquick™ PCR purification kit (QIAGEN). Purified PCR-products were sequenced by GATC Biotech AG (Konstanz, Germany). Both strands were sequenced for new haplotypes. In all other cases (except when sequence chromatograms were of poor quality) only one strand was sequenced. One sequence of every observed haplotype was deposited in GenBank (access. numbers AY 600064-AY 600082).
Data Analysis
An alignment of 69 ITS sequences was calculated with the program ClustalW as implemented in the software BioEdit (Hall 1999) and manually adjusted. In order to display the phylogenetic relationships within the dataset, a haplotype network was calculated under the 95% parsimony probability criterion (Templeton et al. 1992) using the software TCS version 1.06 (Clement et al. 2000) with gaps treated as a 5" character state. Root probabilities (outgroup weights) were calculated for each haplotype according to the heuristic algorithm introduced by Castelloe & Templeton (1994) and according to the modified algorithm by Cassens et al. (2003). Parameters of genetic diversity were calculated using DNASP version 3.99 (Rozas & Rozas 1999). Parameters were calculated for the two European and three Ecuadorian populations from which four or more individuals were sampled (Tab. 2). In addition, genetic diversity within Europe and Latin America respectively was estimated on the basis of all sampled specimens. As estimators of genetic diversity, we calculated haplotype diversity h (Nei 1987) and nucleotide diversity Pi (st). All calculations were performed on two different datasets: one containing all sampled haplotypes and one excluding the deviating European haplotypes H1 and H2 (see below). DNASP ignores positions with gaps, and consequently pools haplotypes that only differ by indels. In order to calculate correct haplotype diversity indices, we recoded positions 12, 13, 18 and 140 of our alignment. Haplotype diversity indices of populations were compared using a two- tailed t-test as described in Zar (1999; equations 8.62 and 8.65) and significance values obtained by an internet application (URL: http://home.clara.net/sisa/signif.htm). The calculation of nucleotide diversity was performed on the original alignments without recoding. Because of multiple hits at positions 69, 90, 113 and 353 of the alignment we used the Jukes and Cantor correction (Jukes & Cantor 1969) to calculate Pi (at). We used an analysis of molecular variance (AMOVA, Excoffier et al. 1992) to partition the total genetic variance of our sample into components due to differences within populations, among populations within continents (Europe and Latin America), and among continents using Arlequin version 2.001 (Schneider et al. 2001). For this analysis we used the same five populations as for the calculation of genetic diversity. Again, calculations were performed on the two datasets outlined above.
Results
The complete aligned dataset is 530 bp long and contains 48 polymorphic sites. Nineteen haplotypes can be distinguished (Tab. 1). The cladogram in Fig. | consists of two clades that are not connected to each other under the 95% parsimony probability criterion. The two haplotypes H1 and H2 are separated from the rest of the network by at least 21 mutational steps. This distance is almost as large as the minimum distance
47
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48
between our two unconnected clades. More specifically, the genetical differences do not coincide with the supposed morphological differences between T. glaucolepidea and T. percrenata. Alternatively, these distant haplotypes could also represent paralogous ITS sequences. In the context of the present study we were unable to check for paralogs and hence conducted all of our analyses on two datasets, one including and one excluding H1 and H2. Excluding H1 and H2 reduces the number of polymorphic sites to 30.
Fig. 1: Haplotype network of 69 ITS sequences from Trapeliopsis glaucolepidea. Ovals represent haplotypes sampled by us. The size of each oval is proportional to the number of sampled individuals of that haplotype. A line represents one mutational step. Black dots represent haplotypes not present in the sample. Haplotypes H1-H7 (grey) occur exclusively in Europe, H8-H19 (white) only in Latin America. The subnetwork consisting of haplotypes H1 and H2 is not connected to the main network under the 95% parsimony probability criterion. The minimum distance between both subnetworks is 21 steps. The black arrow indicates the probable connection between H1 and the large subnetwork. Bold marginal lines indicate alternative rooting positions of the large subnetwork: grey according to the algorithm by Castelloe & Templeton (1994), black according to the modified algorithm by Cassens et al. (2003).
Fig. 1 shows that Latin America and Europe do not share any haplotypes. However, the haplotypes from each continent do not form mutually monophyletic lineages. The European haplotype H7 clusters among the Latin American haplotypes close to the oldest — i.e. most interior — haplotypes of the network. The ancestral haplotype inferred by the algorithm of Castelloe & Templeton (1994) is the Ecuadorian H17 near the periphery of the cladogram. However, it has recently been emphasized that this algorithm is problematic when intermediate haplotypes are missing from the dataset or when the frequency of haplotypes is unequal over the geographical range of a species
49
(Cassens et al. 2003, Printzen et al. 2003). In our case the high outgroup weight of H17 is entirely due to the frequency of its immediate neighbour, H16, in the Ecuadorian population El (see below). With the modified algorithm by Cassens et al. (2003), by far the highest outgroup weight is assigned to H11 in the center of the network, which also occurs in Ecuador.
The genetic diversity indices for European and Latin American populations are summarized in Tab. 2. Two trends are obvious from the table. Firstly, the number of haplotypes is considerably higher in Latin America (12) than in Europe (5 or 7, including H1 and H2). Haplotype numbers differ also between single populations from different continents. Secondly, the inclusion of the deviating haplotypes H1 and H2 grossly inflates the nucleotide diversity but has relatively little effect on haplotype diversity. Although Latin America has a much higher number of haplotypes, the haplotype diversity is not significantly higher than for Europe (Tab. 3). It becomes significant only when H1 and H2 are removed from the European sample. A direct comparison of single Latin American and European populations indicates that only one European population (C3) displays a significantly lower diversity than the Latin American populations (Tab. 3).
Table 2: Number of observed haplotypes and diversity indices for two European and three Ecuadorian populations of 7. glaucolepidea, all Latin American populations and European
populations including and excluding the deviating haplotypes H1 and H2.
Population N__ No. of haplotypes Haplotype diversity (JC) Czech Rep. 1| (excl. H1-2) 12 4 0.636 0.0231 Czech Rep. 1 (excl. H1-2) 5 3 0.700 0.0015 Czech Rep. 3 2 0.286 0.0011 Ecuador 1 17 6 0.779 0.0015 Ecuador 2 3] 0.857 0.0022 Ecuador 3 4 4 1.000 0.0038 Europe (incl. H1-2) 38 7 0.781 0.0205 Europe (excl. H1-2) Dah, 5) 0.689 0.0096 Latin America BHI 12 0.862 0.0036
The fact that no haplotypes are shared between Europe and Latin America is reflected by the high percentage of genetic variation between the continents. According to the results of the AMOVA (Tab. 4) all three hierarchical layers (among continents, among populations within continents and among individuals within populations) contribute more or less equally to the total genetic variance of our sample when H1 and H2 are included. When these haplotypes are excluded more than 70% of the total genetic variance is due to differences between the continents.
Morphological differences between the specimens are apparently not associated with genetic differences in the studied gene. Our results corroborate the opinion that T. glaucolepidea and T. percrenata are conspecific (Palice 1999). The ITS-sequences of British samples representing typical T. glaucolepidea are identical to some sequences from small, wood/peat-inhabiting morphotypes from Belgium, the Czech Republic,
50
Germany, Poland and Sweden (“‘typical” T. percrenata).
Table 3: Comparison of haplotype diversity indices and significance of results according to a two-sided f-test. Significant differences (at the 0.05 level) are found between Latin America and Europe excluding H1 and H2, and Czech Rep. 3 and three Ecuadorian populations.
Population Latin America Ecuador 1 Ecuador 2 Ecuador 3 Europe (incl. H1-2) 0.14 - - = Europe (excl. H1-2) 0.02 * - - - Czech Rep. | (incl. H1-2) - 0.34 O25 0.13 Czech Rep. 1 (excl. H1-2) - 0.74 0.56 0.31 Czech Rep. 3 - 0.04 * G03" 0.02% Discussion
The fact that there are no shared haplotypes among the two continents and the results of the AMOVA provide strong evidence against recurrent genetic exchange between Latin American and European populations of T: glaucolepidea. This is in line with the results of Printzen et al. (2003) who concluded that there is no evidence for recurrent gene flow between Western and Eastern North American and European populations of Cavernularia hultenii. However, it is still controversial, whether rare dispersal events in the distant past can account for the wide disjunctions displayed by many lichens. Recently, Hégberg et al. (2002) reported evidence for a North American origin of European populations of Letharia vulpina. Their inference of long range dispersal is based on the fact that there is higher genetic variability in North America than is observed in European populations of L. vulpina. According to Hégberg et al. (2002) the low diversity in European populations is best explained by a historical population bottleneck that was in turn the results of a founder event after long range dispersal from North America to Europe. A similar result — relatively low genetic diversity in European as compared to American populations — was reported by Printzen et al. (2003) and is also found here. We hesitate, however, to infer historical long range dispersal on the basis of these data. Low genetic variability in certain populations can be explained by population bottlenecks (Nei et al. 1975). However, there may be alternative explanations to a founder event through long range dispersal for such a bottleneck. Most boreal and temperate species of the Northern Hemisphere underwent pronounced population bottlenecks and changes of genetic diversity during the Pleistocene glaciations (e.g. Hewitt 1996, 1999, Soltis et al. 1997, Taberlet et al. 1998). These bottlenecks could have been more pronounced in Europe with its east-west extending mountain ranges than in North America. Their effects on tropical Latin American populations will certainly have been different from those on temperate European populations. The fact that European and Latin American haplotypes of 7. glaucolepidea do not even form monophyletic lineages, would make it necessary to assume at least two independent events of historical long range dispersal, or to assume a very ancient dispersal event with subsequent lineage sorting between the continents. In our opinion, the data currently available on the genetic diversity of lichens is by far too poor which makes such assumptions largely speculative.
51
Table 4: Results of the AMOVA including the five populations Czech Rep. 1-2 and Ecuador 1-3 based on the total dataset and the dataset excluding haplotypes H1 and H2 (in bold).
ae Variance Percentage of Source of variation d. f. Sum of squares ‘ is components variation
Is 1 77.421 2.32643 Va 32.58
wrote P edad 73.793 4.07732 Va 73.16
Among populations within 3 SOM 2 1.98713 Vb 27.62
groups 3 14.338 0.55365 Vb 9,96
Withi at 41 117.179 2.85801 Vc 39.73
ary ale nieth resviry 34 31.079 0.91408 Ve 16.45 Total 45 250.196 7.19361 i 38 118.667 5.55712
It appears from our data that the number of haplotypes is higher in the tropics than in Europe. There is also a trend to higher haplotype diversity, however, direct comparisons between European and Latin American populations are mostly insignificant. These results reflect the fact that most Latin American haplotypes have only been collected once and should not be overinterpreted. Most of our tropical samples originate from only a small area. If some of the rare haplotypes are more common in other tropical areas, a more comprehensive sampling and sequencing effort would probably lead to a higher haplotype diversity in the tropics as could be expected from the number of haplotypes alone.
Nucleotide diversity Pi (st), however, is not necessarily affected in the same way. The pairwise, average distance between sequences depends largely on the distance between the most common haplotypes in a sample. In our case, the genetic distance between the most common European haplotypes (H1,) H4 and H7 is much larger than the distance between any of the Latin American haplotypes. We therefore conclude that nucleotide diversity is not a useful measure of genetic diversity, when comparing populations from different regions with non-overlapping haplotype composition.
Because closely related lichen species may be difficult to distinguish morphologically, species delimitations are often controversial in lichenology. For example, while some recent floras and checklists treat T. glaucolepidea and T. percrenata as different (Foucard 2001, Purvis et al. 1992, Sgchting & Alstrup 2002), others suggest that both are identical (Hafellner & Tiirk 2001, Santesson et al. 2004, Vézda & LisSka 1999). If morphological differences between species were associated with genetical differences, this would support the presence of two species instead of one. However, in our present dataset, morphologically different individuals often belong to the same haplotype, and morphologically similar ones are spread over the haplotype network.
A related question that has repeatedly been discussed in recent times is whether genetically differentiated populations of lichens form cryptic species (Grube & Kroken 2000, Kroken & Taylor 2001). The fact that European and Latin American populations are apparently isolated from each other could indicate the presence of two cryptic species. However, we caution against the over-use of the term “cryptic species” for every instance of genetic differentiation. Reproductive isolation may be a prerequisite
a)
for the formation of species but does not, in itself, constitute speciation. More genetical data from a broader range of markers and a more complete sampling over a wider geographical range would be necessary to assess the species status of temperate and tropical populations of T. glaucolepidea. At present, there is no evidence for cryptic speciation in our dataset.
Acknowledgements
The authors wish to thank Martin Grube (Graz) and Thorsten Lumbsch (Chicago) for a critical review of the manuscript. André Aptroot (Baarn), Blanka Buryova (Pruhonice), Pawel Czarnota (Poreba Wielka), Josef Halda (Rychnov n. Kn.), Harrie Sipman (Berlin), Ondrej Peksa (Praha) and Pieter van den Boom (Son) are thanked for kindly providing lichen samples. The staff of the Experimental department of the Botanical Institute in Pruhonice (Judith Fehrer, Lida Kirschnerova, Maria Loncova, Marie Stara) is warmly appreciated by ZP for their versatile help in the lab. The work was supported by grants of the Grant Agency of the Academy of Sciences of the Czech Republic: KJB6005306, KSK6005114, and AVOZ6005908, and the Grant Agency of the Czech Republic: 206/98/1194.
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Zar JH. 1999. Biostatistical Analysis. 4° ed. New Jersey, Prentice Hall.
54
Appendix 1: Populations sampled for this study
Country
Scotland Ireland Netherlands Belgium Germany Germany Germany Sweden Lithuania Poland Poland Poland
Poland Czech Rep. - C1
Czech Rep. - C2 Czech Rep. - C3
Czech Rep. - C4
Ecuador - El
Ecuador - E2 Ecuador - E3 Ecuador - E4 Ecuador - E5
Costa Rica
Haplotype(s)
H7 7 H5 H7 H1 H7 H7 Hy H1 H2 H1 H7
H7
H1, H4, H5, H6
H7 H4, H5
H3;,H7
H9 -H10, Hil; H16, H17, H19
HUTS 416,
Hi 7 irs H12,H13,H14,
H15 H15
H16
H8
Coordinates
(WGS 84)
56°40-45’N, 03°34-44’W 51°48°N, 09°27°W 52°15’N, 05°40’E 50°27-30’N, 06°06’ E 48°55°N, 13°25°E 52° 15°N, 14°21E 50°06’N, 06°40°E 55°54’N, 13°33°E 54°32’N, 23°49°E 53°40’N, 18°15°E 53°33°N, 17°43’E 50°49’N, 15°22°E 49°10’N, 20°05’E 48°41-55°N, 13°50’- 14°03’30"E 49°05’N, 13°31-34’E 50°53-54°N, 14°23-25’E 49°38-40’N,
15°53’-16°04’E
00°48-49’N, 77°55-S7T W
01°07-17’S, 78°19-21°W 04°08-11S, 79°06-10’W 01°41’S, 78°27 W 00°18’S, 78°12°W 09°34°N, 83°45°W
Collector (Herbarium)
B.Buryova (hb. Palice 6962,
6975) : . J.Halda & Z.Palice (hb. Palice 8063b)
A.Aptroot 48260 (hb. Palice) B. & P. v.d.Boom 28370, 31546 (hb. Palice)
J.Halda et al. (hb. Palice 2672) B. & P. v.d.Boom 28298 & H.Sipman (hb. v.d.Boom) A.Aptroot 59313 (hb. Palice)
Z.Palice (hb. Palice 8201) P.Czarnota (GPN/3117) P.Czarnota (GPN/2961)
P.Czarnota (GPN/3205)
P.Czarnota (hb. Palice, dupl. ex GPN/3345)
P.Czarnota (GPN/2894)
Z.Palice (hb. Palice 1481, 1601, 4212, 4850, 5094, 5101, 6818,
7036, 8630, 8631, 8632, 8633) Z.Palice, O.Peksa (hb. Palice
6551, 8629) Z.Palice (hb. Palice 5808, 5814,
5831, 6832, 6834, 8605, 8606) Z.Palice & O.Peksa (hb. Palice
8066, 8067)
Z.Palice, (hb, Palice,23.12.,2323, 3065a, 3065b, 3255, 8494, 8495, 8496, 8497, 8499, 8503, 8505, 8507, 8508, 8509, 8515, 8519) Z.Palice (hb. Palice 4336, 4348, 8504, 8511, 8512, 8513, 8524) Z.Palice (hb. Palice 2707, 3149,
3578, 3781) Z.Palice (hb. Palice 3203)
Z.Palice (hb. Palice 2234)
H.Sipman 11792 (B)
1
1
MYCOTAZXON
Volume 90(1), pp. 55-68 July-September 2004
New and rare rust fungi (Uredinales) from Anatolia (Turkey) - 2
ZELIHA BAHCECIOGLU zbahcecioglu@inonu.edu.tr
Inonii University, Pharmacy Fac. Malatya, Turkey
HALvor B. GJAERUM
plantevernet@planteforsk.no Norwegian Crop Research Institute, H¢gskoleveien 7, N-1432 As, Norway
Abstract—Twenty-two rust taxa are reported from Anatolia in Turkey. Three of them are described as new species, Phragmidium sarcopoterii on Sarcopoterium spinosum, Puccinia onosmaticola on Onosma molle and Uromyces dorystoechadis on Dorystoechas hastata. Puccinia tepperi on Phragmites australis is reported for the first time outside Australia while four other taxa are reported for the first time in Turkey: Ochropsora ariae on Anemone blanda, Frommeélla tormentillae on Potentilla sp. and Puccinia constricta on Teucrium orientale. Some new hosts for previously known species in Turkey are reported.
Key words—microfungi, new taxa, parasite
Introduction
As Bahcecioglu & Gjzerum (2003) have previously observed, Turkish mycota is rich in microfungi. Some research has been done, e.g. by Karel (1958) and Henderson (1959, 1961, 1964), but there is still much to do.
Materials and methods
The material presented here was collected in the years 1995-2002 in the following provinces: Agri, Anamur, Antalya, Bitlis, Erzincan, Erzurum, Giimtishane, Kahramanmaras, Malatya, Ordu and Sivas. Spores were mounted in lactophenol. The host names follow the Flora of Turkey and Aegean Islands by Davis (1968 — 1985) and Davis et al. (1988). After identification of the rusts the material is preserved in the herbarium of Indnii University.
56
Frommeélla tormentillae (Fuckel) Cummins & Y. Hirats. — Illustrated genera of rust fungi. Rev. ed., p. 120, 1983.
Syn. Frommea obtusa (Fuckel) Arthur. — Bull. Torrey Bot. Club 44: 503, 1917. On Potentilla sp. (Rosaceae).
Giimiishane. Zigana Pass, 1950-2000 m, 12. Aug. 2001, Z. Bahcecioglu 3228, II+III.
Uredinia hypophyllous, pale yellow with a few clavate, thin-walled paraphyses. Urediniospores 19-22 x 16-20 ym, subgloboid, ellipsoid or obovoid, walls 1 (-1.5) wm thick, hyaline, finely echinulate with 3-4 indistinct pores. Telia hypophyllous, brown. Teliospores 54-84 x 27-32 wm, cylindric or clavate, 3-6-celled, mostly 5, with one pore in each cell, walls brown, smooth, at side 4-5 yum thick, apically thickened to 4.5-6.5 wm, mostly rounded, but often with a short papilla. Pedicels persistent, equal to the length of the spores, hyaline but tinted near the spore, collapsing.
As Frommea obtusa, this rust has been reported on many species of Potentilla in Europe and the Americas, more rarely in Africa and Australia. In Asia it is reported from Armenia, India and Siberia. Frommeéila is a new genus in the Turkish rust flora.
Ochropsora ariae (Fuckel) Ramsb. — Trans. Brit. Mycol. Soc. 4: 337, 1915.
On Anemone blanda Schott & Kotschy. (Ranunculaceae). Malatya. Beydagi. Birticek hill, 2000 m, 01. May. 1995, Z. Bahcecioglu 179, 0+I.
Pycnia epiphyllous, type 7. Aecia amphigenous, mostly hypophyllous, covering the whole leaf, cupulate, peridium white. Aeciospores in chains, 14- 20(-23) x 14-17 pm, ellipsoid or globoid, often somewhat angular, walls c 1 pm thick, densely and finely verrucose, hyaline.
This rust species, systemic in the rootstocks and host alternating with species of Sorbus, occurs on many species of Anemone from Europe to China and Japan. Woronow (1910) and Durrieu (1968) reported A. blanda as a host for this rust from Caucasus and Greece, respectively, both as O. sorbi (Oud.) Dietel. Ochropsora is a new genus to the rust flora of Turkey.
Phragmidium sarcopoterii Gjz2rum & Bahcecioglu, sp. nov. (Fig. 1).
Etymology: from Sarcopoterium, a genus in Rosaceae.
Pycnia, aecia et uredinia ignota. Telia hypophylla, atrofusca cum paraphysibus clavatis vel cylindricis, pariete 1-1.5 jm crasso, hyalino. Teliosporae 47-80 x 31-39 wm, (3-)4-5(-6)- cellularis, cylindraceae, ad basem leviter dilatatae, ad apicem rotundatae, papillis hyalino vel apiculato verrucoso usque ad 15 wm longo, pariete 4-4.5 wm crasso, ad apicem 4.5-5.5ym incrassato, verrucis disperso, hyalino, quaque cellula poris duobus, cellulae apicalis raro
S/
tribus instructis, pedicello persistente, usque ad 90 wm longo, vulgo deorsum incrassato, hyalino.
Pycnia, aecia and uredia not seen. Telia hypophyllous, blackish brown with clavate to curved paraphyses, walls hyaline, 1-1.5 wm thick. Teliospores 47-80 x 31-39 ym, (3-)4-5(-6)-celled, cylindric, slightly wider at base, apically rounded with a low, hyaline papilla or often with a warted, abrupt apiculus up to 15 wm long, walls 4-4.5 wm thick, brown, apically thickened to 4.5-5.5 wm, with scattered, hyaline warts, each cell with 2 pores, apical cells rarely with 3 pores. Pedicels up to 90 ym long, generally swelling at base, hyaline.
Holotype: Z. Bahcecioglu 2674 (Indnii), Turkey, Anamur, Between Anamur and Gazipasa, Kaledran village, 50 m, 28. May. 2000, on Sarcopoterium spinosum (L.) Spach. (Rosaceae). Isotype in NCRI.
Fig. 1. Phragmidium sarcopoterii . Teliospores and paraphyses from type.
The teliospores in the present specimen differ from those in Phr. sanguisorbae (DC.) J. Schrét. ssp. mediterranea D. M. Henders. in having thicher walls with scattered warts and often an apiculus and with pedicels longer than the spores.
Phr. sanguineum (DC.) J. Schrot. has been reported on the present host from Libya, Greece and Israel. The ssp. mediterranea is reported from Spain to Greece.
Puccinia calcitrapae DC. — FI. Fr. 2: 221, 1805.
Syn. P. echinopis DC. — FI. Fr. 5: 57, 1815.
On Echinops galeticus Freyn. (Asteraceae).
Sivas. Imranli, Saruhan village, 2000 m, 16.Aug. 2002, Z. Bahcecioglu 3352, (II+) II. On Echinops melitenensis Hedge & Hub.- Mor.
Sivas. Giirtin, Gékpinar, 1400 m, 16. Aug. 2002, Z. Bahcecioglu 3326, (II+) III.
Uredinia not seen, but some spores have been found in the telia. Urediniospores 26-34 x 26-31 ym, ellipsoid or subgloboid, walls 1.5-3 ym
58
thick, slightly thickened basically, yellowish brown, echinulate with 3 equatorial pores. Telia amphigenous, mostly hypophyllous, black, small, round, often in rings around a central sorus. Teliospores 33-43(-46) x 23-27 wm, cylindrical, ellipsoid or obovoid, walls 3 wm thick, apically rarely thickened up to 4.5-5 pm, apically rounded, flattened or attenuate, verruculose, brown, upper pore apical or subapical, lower pore somewhat depressed. Pedicels short, hyaline.
P. echinopis is widespread in Central and South Europe through Asia to China, but not in more tropical areas. In Africa it is reported from Morocco, Algeria and Sudan. From Turkey it has been reported as P. echinopis on many species of Echinops as E. heldreichii Boiss., E. aff. viscosus DC. (Henderson 1961) and E. melitenensis Hedge & Hub.- Mor. and E. purgans Trautv., E. ritro L. (Bahcecioglu & Isiloglu 1996, Bahcecioglu & Yildiz 2001). For other asteraceous genera serving as hosts for this rust see e.g. Henderson (1964). P. echinopis 1s now recognized as a race or a f. sp. of P. calcitrapae s. lat.
Puccinia caricina DC. s. lat. — Fr. Fl. 5: 60, 1815. On Carex sp. (Cyperaceae). Ordu. Between Ordu and Aybasti, 1500-1600 m, 17. Aug. 2001, Z. Bahcecioglu 3325, II+III.
Uredinia hypophyllous, oblong, often confluent, covered by epidermis which later cracks, brown. Urediniospores 27-33 x 17-25 vm, subgloboid, ovoid or obovoid, walls 1.5-2.5 ym thick, yellow, echinulate, with 3 equatorial pores, covered by low, hyaline papillae. Telia mainly hypophyllous, oblong, often confluent, covered by epidermis which later cracks, dark brown. Teliospores 47-65 x 15-22 ym, clavate, slightly constricted, apically rounded or obtuse, walls brown, in lower cells about 1 wm thick, in upper cells 1.5 ym thick, apically thickend up to 10 wm, smooth, upper pore apical, lower near septum. Pedicels persistent, pale brown, up to 25 ym long.
P. caricina is reported from all continents, common on many species of Carex in Europe, but also in Asia and in the Americas, more rarely in Africa and Australia. The first report of this rust from Turkey was given by Magnus (1903) who reported it as P. caricis (Schum.) Rebent. on C. hirta L. Later Henderson (1964) and Bahcecioglu & Gjerum (2003) reported it on C. hordeistichos Vill., Henderson also on C. sylvatica Huds.
Puccinia chaerophylli Purton. — Midland Flora 3: 303, 1821. On Chaerophyllum macrospermum (Sprengel) Fisch. & Mey. (Apiaceae). Giimiishane. Zigana Pass, 1600 m, 12.Aug.2001, Z. Bahcecioglu 3219, II+III.
Uredinia hypophyllous, rounded or oblong, yellowish brown. Urediniospores 21-27 x 20-23 ym, subgloboid or ellipsoid, walls 1.5-2.5 wm thick, pale yellow or yellowish brown, echinulate with 3 equatorial pores
59
covered by low, hyaline papillae. Telia amphigenous, mostly hypophyllous, blackish brown. Teliospores 23-36 x 17-25 ym, ellipsoid or obovoid, often irregular, rounded at both ends or apically obtuse or attenuate below, slightly constricted, walls 2-2.5 wm thick, reticulate, yellowish brown to brown, upper pore apical, lower pore subequatorial. Pedicels hyaline, short.
P. chaerophylli is common in Europe and Asia except in tropical areas, rare in Africa. In Turkey it has previously been reported on Anthriscus nemorosus (Bieb.) Sprengel (Tamer & Oner 1978), Chaerophyllum byzantinum Boiss. (Gjzrum 1996) and C. bulbosum L. (Bahcecioglu & Yildiz 1996). C. macrospermum is a new host for this rust.
Puccinia constricta (Lagerh.) Bubak. — Centralbl. f. Bacteriol., Parasitenk., Infektionskrankh. II. Abt. 9: 919, 1902.
On Teucrium orientale L. (Lamiaceae).
Kahramanmaras. Goksun, Ahmetcik village, 1300 m, 08. Aug. 2001, Z. Bahcecioglu 3187, II.
Telia on leaves and stems, dark brown. Teliospores 39-48 x 20-31 pm, ellipsoid, upper cells mainly slightly wider than the lower ones, slightly constricted, rounded in both ends or apically slightly attenuate, walls 1-1.5 7m thick, in upper cells gradually thickened to 7 wm, brown, in lower cells slightly paler. Pedicels up to 65 ym long, pale brown, collapsing.
Guyot & Massenot (1952) separated the rusts on Teucrium in two groups, one with pale to reddish brown telia mainly on leaves, the other with blackish brown telia mainly on stems, but also on leaves. In the latter group they recognized P. constricta, P. istriaca P. Syd. & Syd. and P. polii Guyot. They found P. majoricensis Maire similar to P. polii which is the younger name of these two, but they kept them under P. polii. Petrak (1953) and J@rstad (1962) used P. majoricensis with P. polii as a synonym.
P. majoricensis has longer teliospores than both P. constricta and P. istriaca which hardly can be separated that way as they overlap. However, P. constricta has pedicels equal to the length of spores while P. istriaca has short pedicels. Our specimen has teliospores whose pedicels are equal or even longer than the spores. We have therefore placed it under P. constricta which is known in Central Europe from France to Romania and Bulgaria and also in Caucasia. It is new to the rust flora of Turkey.
Puccinia hieracii Mart. var. piloselloidarum (Probst) Jorst. — Kgl. Norske Vidensk. Selsk. Skr. 1935, 38: 27, 1936.
On Pilosella x auriculoides (A. F. Lang) Sell & West. (Asteraceae). Erzincan. Kemaliye, Sirakonaklar village, 1300 m, 24. Jun. 2000, Z. Bahceciolglu 2924, II+II. On Pilosella verruculata (Link) Sojak.
Erzincan. Kemaliye, Sirakonaklar village, 1600 m, 24.Jun.2000 m, Z. Bahcecioglu 2936, II+III.
60
Uredinia hypophyllous, brown. Urediniospores 27-31 x 23-29 ym, subgloboid, ovoid or obovoid, walls 1.5-2.5 wm thick with two pores slightly superequatorial, brown, echinulate. Telia hypophyllous, brown. Teliospores 35-46 x 22-27 ym, ellipsoid or obovoid, somewhat irregular, some spores slightly constricted, walls 2-2.5 wm thick, brown, finely verruculose, upper pore subapical, the lower pore equatorial, pedicels hyaline, short, mostly broken near the spore.
P. hieracii, living on species of many asteraceous genera, is a complex species of which some races or forms have been described as species or varieties, all with two pores more or less superequatorial. P. hieracii var. piloselloidarum is bound to the Pilosella and morphologically characterized by Urediniospors with slightly superequatorial germ pores.
P. hieracii is reported from Turkey by e.g. Magnus (1903), Henderson (1961, 1964) and Bahcecioglu and Yildiz (2001) while Tamer et al. (1987) published the variety on Pilosella x auriculoides
Puccinia lapsanae Fuckel. — Jahrb. Nass. Ver. Nat. p. 13, 1860.
On Lapsana communis L. (Asteraceae). Sivas. Kizilda& Pass, 2100 m, 14. Aug. 1996, Z. Bahcecioglu 1134, II+II; Erzincan. Kemaliye, Sirakonaklar village, 1600 m, 24. Jun. 2000, Z. Bahcecioglu 2941,:II+III; Giimiishane. Torul,
between Zigana village and Zigana Pass, 1600 m, 08.12.2001, Z. Bahcecioglu 3225, II+III.
Uredinia and telia amphigenous, brown, small, often confluent. For the specimens 1134 and 2941 urediniospores 22-26 x 16-22 wm, globoid or subgloboid, walls 1-1.5 jm thick, pale brown, verruculose with 2 equatorial pores. Teliospores 27-35 x 22-25(-27) um, ellipsoid or obovoid, walls 1.5 - 2(-2.5) wm thick, brown, upper pore apical, lower equatorial. Pedicels short, hyaline, deciduous.
Fig. 2. Puccinia lapsanae. Telio- and urediniospores from specimen Z. Bahcecioglu 3225.
61
In the specimen 3225 uredinio- and teliospores have slightly thicker walls and the teliospores (Fig. 2) are often more irregular than in the two other specimens studied. Henderson (1964) mentioned that in S. W. Asia teliospore- walls often are slightly thickened compared to the normal, probably correlated with the regional arid conditions. However, for describing the fungus on No. 3225 e. g. as anew variety we think more material is necessary.
P. lapsanae is common in Europe, but seems to be rare in the other continents. Bahcecioglu & Isiloglu (1996) and Bahcecioglu & Yildiz (2001) reported it on L. communis and Jurinea consanguinea L. from the Malatya province.
Puccinia libani Magn. — Verh. Zool.-Bot. Ges. Wien 1: 442, 1900. On Prangos pabularia Lindl. (Apiaceae). Sivas. 32 km from Giiriin to Pinarbas1, 1800 m, 06. Jun. 1997, Z. Bahcecioglu 1686 m, I+III.
Pycnia not seen. Aecia epiphyllous in long groups, peridial cells rhomboid, 23-39 x 15-20 ym, inner walls verrucose, outer walls smooth, both 6-6.5 xm thick. Aeciospores 21-35 x 19-25 wm, ellipsoid or subgloboid, walls less than | wm thick, densely verruculose, hyaline. Telia epiphyllous, round or oblong, often confluent or connected with the aecia, dark brown. Teliospores 56-72 x (23-)28-35 wm, clavoid, somewhat constricted at septum, walls about | pm thick, apically thickened up to 11 ym, apically rounded or truncate, reddish brown or brown, basically yellowish brown, smooth, upper pore apical, lower pore obscure. Pedicels tinted, short. One-celled spores occur.
The rust is known from Greece and Lebanon east to Kazakhstan. Lindroth (1902) made the first record of this rust in Turkey on P. uechtritzii Boiss. & Hausskn. Henderson (1959, 1964) added P. ferulacea Lindl., P. lophoptera Boiss. and also Peucedanum pisidicum Boiss. P. pabularia is a new host for this rust.
Puccinia onosmaticola Gjzrum & Bahcecioglu, sp. nov. (Fig. 3).
Etymology: from Onosma, a genus in Boraginaceae.
Pycnia, aecidia et uredinia ignota. Telia hypophylla, dispersa, parva, pulverulenta, brunnea. Teliosporae 22-34(-39) x 17-22 ym, ellipsoideae, ad basem raro attenuatae, pariete 3-4 wm crasso, laeve, brunneo, poro superiore subapicali, poro inferiore aequatorio instructis, pedicello brevi, pariete tenui, leviter tinto.
Pycnia, aecia and uredinia not seen. Telia hypophyllous, scattered, small, pulverulent, brown. Teliospores 22-34(-39) x 17-22 ym, ellipsoid, rarely obovoid, rounded at both ends or rarely attenuate at the base, walls 3- 4 um thick, smooth, brown, upper pore subapical, lower equatorial. Pedicels
62 short, slightly tinted.
Holotype: Z. Bahcecioglu 2054 (Inénii), Turkey, Sivas, Giiriin, Gdkpinar, 1500 m, 18. Jul. 1998, on Onosma molle DC. (Boraginaceae). Isotype in NCRI.
The teliospores of this species are shorter with evenly thick walls and the lower pore equatorial while teliospores in P. onosmatis Gjerum & Bahcecioglu are thickened apically and with the lower pore near septum.
5 $§$~-rerethstieemnnrnerlriisisinaibannsninensnnn
4Om 2
Fig. 3. Puccinia onosmaticola. Teliospores from type.
Puccinia pulvinata Rabenh. — Hedwigia 10: 20, 1871.
On Echinops ritro L. (Asteraceae). Sivas. Giriin, Sugul valley, 1400 m, 02. Oct. 2002, Z. Bahcecioglu 3426, II+III. On Echinops sp.
Anamur. Between Anamur and Gazipasa, 50 m, 28. May. 2000, Z. Bahcecioglu 2673, (II+)IIL.
Uredinia hypophyllous, yellowish brown. Urediniospores 37-45 x 34- 39 ym, subgloboid, walls 2-3 wm thick, yellowish brown, apically thickened up to 4.5 wm, at the base up to 4 ym, echinulate, 3-4 equatorial pores covered by wide, hyaline papillae. Telia amphigenous, pulverulent, dark brown. Teliospores 50-63 x 31-39 ym, ellipsoid or ovoid, walls 2-4(-4.5) jm thick, apically thickened up to 5.5 wm, brown, paler above the apical pore, lower pore equatorial, very finely verruculose, easiest to see on dry spores. Pedicels at least up to 130 wm long, in the literature given up to 190 wm (Kapsanaki- Gotsi 1986). One-celled spores occur.
P. pulvinata has been reported scattered from Morocco and Algeria east to Iran and Pakistan. From Turkey it has been reported on E. heldreichii Boiss. (Magnus 1903) and on E. viscosus DC. (Henderson 1964). E. ritro seems to be a new host for this rust species. Other Turkish hosts are Jurinea depressa C. A. Mey. var. pinnatiseta Boiss. and J. pontica Hausskn. & Freyn. (Henderson op. cit.) and Acantholepis orientalis Less. (Gjerum 1996).
63 Puccinia recondita Roberge ex Desm. — Bull. Soc. bot. Fr. 4: 798, 1857.
On Cerinthe minor L. (Boraginaceae). Sivas. Taslidere, 1300 m, 03. Jun. 1997, Z. Bahcecioglu 1531, 0+. On Myosatis lithospermifolia (willd.) Hornem. (Boraginaceae).
Sivas. Divrigi, Hébek village, 1300 m, 02.06.1997, Z. Bahcecioglu 1484, O+1.
Pycnia epiphyllous, type 4, and aecia in groups around the pycnia, peridia white, peridial cells 35 x 10-20 ym, outer walls 8-10 ym thick, striate, inner walls 2-3 wm, verrucose. Aeciospores 25-33 x 21-29 ym, subgloboid, ovoid or ellipsoid, often somewhat angular, walls hyaline, c 1 wm thick, finely verrucose, pores scattered, obscure.
These aecial stages most likely belong to the aggregate species P. recondita which has its aecial stage on species of different genera of Boraginaceae, Balsaminaceae, Hydrophyllaceae and Ranunculaceae. From Turkey it has been reported as Aecidium asperifolii Pers. under P. rubigo- vera (DC.) Winter on Nonnea macrocarpa Boiss. & Heldr. and on Anchusa angustissima C. Koch by Magnus (1891) and as P. agropyri Ell. & Ev. on Clematis cirrhosa L. by Bremer et al. (1952). Henderson (1959, 1964) reported it as A. asperifolii on Moltkia caerulea (Willd.) Lehm. and under P. recondita on Clematis orientalis L. If belonging to P. recondita s. lat. Cerinthe and Myosotis are new host genera for this rust in Turkey.
Puccinia serpylli Lindr. — Acta Soc. Fauna Flora Fennica 26: 10, 1904.
On Thymus fallax Fisch. & Mey. (Lamiaceae). Agri. Eleskirt, Tahir mountain, 2400 m, 21.Jul..2000, B. Yildiz 14853, III; Bitlis. Tatvan, Nemrut mountain, 1750 m, 17.Jul. 2000, B. Yildiz 14802 , III.
On Thymus fedtschenkoi Ronninger. Bitlis. Tatvan, Nemrut mountain, 1750 m, 17. Jul. 2000, B. Yildiz 14796, III.
On Thymus kotschyanus Boiss. & Hohen. Bitlis. Tatvan, Nemrut mountain, 1750 m, 17. Jul. 2000, B. Yildiz 14801, III.
Telia hypophyllous, blackish brown, round or ellipsoid, scattered or in small groups, pulverulent. Teliospores 39-53 x 22-29 yum, ellipsoid or obovoid, mostly rounded apically, constricted at septum, walls 1-1.5 wm thick, apically thickened up to 8 ym, smooth, brown. Pedicels up to 115 vm long, thin-walled, hyaline but tinted near the spore, persistent.
P. serpylliis microcyclic. It has been found scattered on a few species of Thymus from Finland and Kola in Russia to Mongolia. Henderson (1959) reported it on P. kotschyanus from Turkey, and it has also been reported on the same host from Iran (Petrak 1953). 7. fallax is a new host for the rust.
Henderson (op. cit.) pointed out the similarity to P. stipina Tranz. f. sp. thymi-stipae Kleb. and indicated it might has been derived from this longcyclic species, alternating between Thymus (0+I) and Stipa (II+HI).
64
Puccinia stipae Arthur var. stipina (Tranzschel) H. C. Green & Cummins. — Mycologia 50: 21, 1958.
Syn. Puccinia stipina Tranzschel — Trav. Mus. Bot. Acad. Imp. Sci. St. Petersb. 7: 114, 1909 (nom. nud). On Thymus kotschyanus Boiss. & Hohen. (Lamiaceae).
Malatya. Arapkir road, Karayaollari bakimevi, 26. Jun. 1999, 1100 m, B. Yildiz 14356, I.
Pycnia amphigenous, type 4. Aecia hypophyllous, in dense groups. Peridia white, long, peridial cells polygonal, 20-40 x 10-12 jm, outer walls up to 8 ym thick, striate, inner walls up to 2 wm, verrucose. Aeciospores 21-27 x 16-19 pm, subgloboid, oblong or ellipsoid, often irregular, walls 1.5-2.5(-4) pm thick, sometimes unevenly thickened, verrucolose, yellowish brown, pores obscure.
P. menthae Pers. has also been reported with aecia on Thymus, but its aeciospore-walls are hyaline.
This taxon is host-alternating between species of Stipa (II+III]) and species of many lamiaceous genera. On Thymus spp. the aecial stage is fairly common in Central and South Europe, but it seems to be very rare in Asia, reported only from Kazakhstan. Tamer et al. (1990) and Giiven & Tamer (1993) published it from Turkey. 7: kotschyanus is a new host for this taxon.
Puccinia tepperi F. Ludw. — Zeitschr. Pflanzenkr. 2: 132, 1892. On Phragmites australis (Cav.) Trin. ex Steudel. (Poaceae). Malatya. 15 km from Malatya to Sivas, 900 m, 19. Jun. 2002, Z. Bahcecioglu 3332, III.
Telia amphigenous, confluent, densely covering large areas of the leaves, brown. Teliospores 43-70 x 21-26(-30) wm, obovoid, ellipsoid or more rarely fusiform, slightly constricted, walls 2.5-4.5 wm, apically thickened to 8.5 wm, yellowish brown, apically paler, smooth, upper pore apical, the lower near septum. Pedicels persistent, thick-walled, not collapsing, up to 220 wm long. It differs from P. moriokaensis S. Ito and P. invenusta P. Syd. & Syd. on the same host, respectively, wider and longer teliospores.
P. tepperi is macroscopically separated from other species of Puccinia on Phragmites as the teliospores are confluent in large, compact groups adaxially on the leaves.
This fungus is so far known only from to the type locality, Grange in South Australia. Cummins (1971) when examining the type (S), found two urediniospores. We have seen none.
Trachyspora intrusa (Grev.) Arthur. — Manual of the rusts in United States and Canada, p. 97, 1934.
On Alchemilla barbatiflora Juzep. (Rosaceae).
Giimiishane. Zigana Pass, 1600 m, 12. Aug. 2001, Z. Bahcecioglu 3215, III.
On Alchemilla mollis (Buser) Rothm.
Sivas. Serefiye, Armutcayiri village, 1500 m, 16. Aug. 2002, Z. Bahcecioglu 3343, III.
65
Telia hypophyllous, sori small, surrounded by the epidermis, often confluent, brown. Teliospores 23-35(-43 ) x (18-)22-27 wm, globoid, obovoid or oblong, walls 2-2.5 wm thick, upper part coarsely verrucose, lower part smooth or nearly so, brown. Pedicels hyaline, 1-septate, upper cells shorter than the lower ones, deciduous.
This rust is widespread in the northern hemisphere. From Turkey it has previously been reported as T: alchemillae (Pers.) Fuckel on A. acutiloba Stev. by Maire (1906) and as Uromyces alchemillae (Pers.) Fuckel on A.. vulgaris L. by Magnus (1891). As A. intrusa it was published by Henderson (1964) on A. vulgaris L. agg., by Tamer et al. (1989) on A. pseudo-castalini Juzep. and Bahcecioglu (2004) on A. holocycla Rothm.
Uromyces dorystoechadis Gjzrum & Bahcecioglu, sp. nov. (Fig. 4).
Etymology: from Dorystoechas, a genus in Lamiaceae.
Pycnia, aecidia et uredinia ignota. Telia hypophylla, parva, dispersa, fusca. Teliosporae 28-31 x 27-30 wm, globoideae, subgloboideae, ellipsoideae vel globoideae-angulares, pariete 3.5-4 wm crasso, fusco, laeve, poro apicalis. Pedicella usque ad 240 ym longi, hyalini.
Pycnia, aecia and uredinia not seen. Telia hypophyllous, small, scattered, dark brown. Teliospores 28-31 x 27-30 vm, globoid, subgloboid or ellipsoid, rarely globoid-angular. Walls evenly 3.5-4 wm thick, dark brown, smooth, pore apical. Pedicels up to 240 wm long, hyaline.
Holotype: Zeliha Bahcecioglu 2892b (Indnii), Turkey, Antalya, Kemer — Goymiik, Degirmendere, 50 m, 29. May. 2000, on Dorystoechas hastata Boiss. & Heldr. ex Benth. (Lamiaceae). Isotype in NCRI.
Fig. 4. Uromyces dorystoechadis Teliospores from typ.
To our knowledge no rust has previously been described on this genus. It differs from other Uromyces species on lamiaceous hosts having teliospores with evenly thickened walls, the others are thickened apically.
66
Uromyces fallens (Arthur) Kern ex Barth. — Handb. N. Amer. Ured. p. 61,1928.
On Trifolium pratense L. (Fabaceae).
Giimiishane. Zigana Pass, 1600 m, 12. Aug. .2001, Z. Bahcecioglu 3221a, II.
Uredinia amphigenous, round, coalescent, brown. Urediniospores (17-)22-30 x 20-24 wm, subgloboid, ellipsoid, obovoid, often somewhat irregular, walls (1.5-)2-2.5 thick, yellowish brown, 3-5(-6) pores scattered or 2-4 equatorial and one apical, echinulate.
This rust is widespread, occurring on many species of Trifolium, but seems to be new to the rust flora of Turkey. Cummins (1977) recognized it as a variety of Uromyces trifolii-repentis Liro, U. trifolii-repentis var. fallens (Arthur) Cummins.
Uromyces punctatus J. Schrot. — Abh. Schles. Ges. Vaterl. Cult. Nat. Abh. 1869-72, 1870.
On Astragalus densifolius Lam. (Fabaceae).
Erzincan. Kemaliye, Sirakonaklar village, 1600 m, 24. Jun. .2000, Z. Bahcecioglu 2942, III.
Uredinia mostly epiphyllous, round or oblong, on stems in long streaks, pale brown. Urediniospores 22-29 x 20-23 vm, obovoid, subgloboid, often angular, walls 1.5-2 wm thick, pale brown with 3-4 pores , when 3 often equatorial, with low, hyaline papillae, echinulate. Telia as the uredinia, but blackish brown. Teliospores 21-28 x 17-20 ym, ellipsoid, obovoid or somewhat irregular, walls 1-1.5 jm, dark brown, verrucose, verrucae often in lines, pore apical, often with a low, hyaline papilla. Pedicels short, hyaline, deciduous.
U. punctatus has been recorded on more than one hundred species of Astragalus (Gjerum 1991) and occurs also on species of Oxytropis. A. densifolius is a new host for this taxon which is often recognized as a race of U. pisi (DC.) Otth (cfr. Henderson 1964). Henderson (1961) reported this rust on A. gummifer Lab. and A. sp. from Turkey.
Uromyces scleranthi Rostr. — Bot. Tidsskr. 21: 40, 1897.
Syn. Urom. alsines Tranzschel. — Ann. Mycol. 5: 547, 1907. On Minuartia hamata (Hausskn.) Mattf. (Caryophyllaceae).
Malatya. 10 km from Malatya to Arapkir, 1600 m, 24. Jun. 2000, Z. Bahcecioglu 2910, III.
Telia hypophyllous, blackish brown. Teliospores 20-25 x 15-21 ym, subgloboid, ellipsoid or obovoid, often somewhat irregular, walls 1.5-2 wm thick, pale brown, apically slightly thickened with a wide, hyaline cap over the apical pore, finely verrucose. Pedicels hyaline, deciduous.
The size of the teliospores in this specimen corresponds fairly well to what is given in the diagnosis, but they have slightly thinner walls.
67
On species of Minuartia (Alsine) this rust has been reported from Bulgaria, Russia (Crimea) and Iran. Bahcecioglu (2001) reported it from Turkey on M. meyeri (Boiss.) Born. and M. hamata. On Scleranthus spp. it has been reported from Denmark, France, Hungary, Russia and Australia.
Zaghouania phillyreae Pat. — Bull. Soc. Myc. Fr. 17: 187, 1901. On Phillyrea latifolia L. (Oleaceae). Antalya. 3 km from Antalya to Gebiz, 100 m, 22. May. 2000, Z. Bahcecioglu 2687, I.
Pycnia not seen. Aecia on stems, amphigenous on malformed, thickened leaves, pustulate, covered by epidermis, orange, peridium short. Aeciospores 18-25 x 15-19 wm, subgloboid or ellipsoid, walls 2-2.5 wm thick, hyaline, alveolate-reticulate.
Z. phillyreae has been reported on Phillyrea in South England (cult.) and in the Mediterranean from Morocco, Algeria and Portugal to Israel. On species of Osmanthus it has been reported from China and Japan. Zaghouania is a new genus in the Turkish rust flora.
Acknowledgements
We are indebted to Prof. Dr. Bayram Yildiz, Balikesir University, Balikesir, for allowing us to examine and publish his specimens of Thymus and for help with identification of the hosts. We also want to thank Eng. Vidar Stensrud, As, for help with preparing the manuscript.
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Gjerum HB. 1991. Studies in rusts (Uredinales) on Astragalus (Fabaceae). Edinb. J. Bot. 48: 393-401.
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Giiven K, Tamer A.U. 1993. Some Parasitic Fungi Determined in Plants Living in Eskisehir. — J. Fac. Sci. Ege University, ser. B, 15: 28-32.
Guyot AL, Massenot M. 1952. Les rouilles des Teucrium. — Bull. soc. bot. suisse 62: 429-475.
Henderson DM. 1959. Uredinales from S. W. Asia. — Notes R. Bot. Gard. Edinburgh 23: 71-83.
Henderson DM. 1961. Uredinales from S. W. Asia: II. — Notes R. Bot. Gard. Edinburgh 24: 249- 258.
Henderson DM. 1964. Uredinales from S. W. Asia: III. The rust fungi of Turkey. — Notes R. Bot. Gard. Edinburgh 25: 197-277.
Jgrstad I. 1962. Investigations on the Uredinales and other parasitic fungi in Mallorca and Menorka. — Skr. Norske Vidensk.- Akad. Oslo. I. Mat.-Nat. Kl. Ny Ser. No. 2: 1-73.
Karaboz I, Oner M. 1982. Parasitic fungi from the province of Manisa. -Mycopathologia 79: 129-131.
Kapsanaki-Gotsi 0. 1986. Contribution to the knowledge of the mycoflora of Kriti island (Hellas). Taxonomic and floristic study on the Uredinales. PhD Thesis. University of Athens. Athens, pp.256 (In Greek).
Karel G 1958. A Preliminary List of Plant Diseases in Turkey, Ayyildiz Matbaasi Ankara.
Lindroth J I. 1902. Die Umbelliferen-Uredineen. — Acta Soc. Fauna Flora Fennica 22: 2-225.
Magnus P. 1891. Ein kleiner Beitrag zur Kenntnis der parasitischen Pilze Kleinasiens. — Engl. Bot. Jahrb. 14: 485-494 + 1 PI.
Magnus P. 1903. Fungi. Ein weiterer Beitrag zur Kenntniss der Pilze des Orients. — In J. Bornmiiller, Iter Anatolicum Tertium 1899. — Bull. l’Herb. Boissier, Sec. sér. No. 7: 573-587 + 2 Pl.
Maire R.1906. Etudes des champignons récoltés en Asie Mineure (1904). —Bull. Soc. Sci. Nancy pp. 165-188. (Reprint pag. 1-26.)
Petrak F. 1953. Beitrage zur Kenntnis der Pilzflora Irans. — Sydowia 7: 50-78.
Tamer A.U, Altan Y, Giicin F. 1989. Giilveren K6yii (Erzurum —Senkaya). Florasinda Belirlenen Bazi Parazit Funguslar. — A. U. Fen Edebiyat Dergesi 1: 45-55. (In Turkish).
Tamer A.U, Altan Y, Giicin F. 1990. Dogu Anadolu Florasinda Belirlenen Bazi Parazit Funguslar. — Turkish J. of Botany 83 — 86. (In Turkish).
Tamer AU, Giicin F, Altan Y. 1987. Malatya (Piitiige), YOresi Bitkilerinde Belirlenen Bazi Parazit Funguslar. — 8. Ulusal Biyoloji Kongresi, Ege University II. 202 - 217. (In Turkish).
Tamer A.U, Oner M. 1978. The parasitic fungi of Aydin Province. —Mycopathologia 64: 87-90.
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MYCOTAZON
Volume 90, pp. 69-76 July-September 2004
A new species of Pseudobaeospora from California
DENNIS E. DESJARDIN
ded@sfsu.edu Department of Biology, San Francisco State University 1600 Holloway Ave., San Francisco, CA 94132 USA
Abstract — Pseudobaeospora stevensii sp. nov. is described from cypress and redwood forests in northern California. Diagnostic features include dark brown basidiomes lacking violet colors, conspicuous cheilocystidia, clamp connections, and tissues that turn olive to grass green or bluish green in alkaline solution. Notes on type studies of several temperate North American species referable to Pseudobaeospora are provided.
Key words — agarics, Basidiomycetes, taxonomy Introduction
The genus Pseudobaeospora was established by Singer (1942, 1963) to accommodate Baeospora oligophylla Singer, a small, violet species described from the Altai Mountains of central Asia (Singer 1938). Since then, numerous species have been described in or transferred to Pseudobaeospora. The generic circumscription was redefined recently by Bas (2002, 2003), wherein he provided a key, full descriptions and illustrations of all known European species. Very few species attributable to Pseudobaeospora have been described or reported from temperate North America, and a critical study of North American taxa has not been published. It should be noted that all reported North American Pseudobaeospora form violet-colored basidiomes that do not discolor (red, green, blue or yellow-green) in KOH solution.
Recently, an undescribed species with basidiomes that lack violet or purple colors and that turn olive to grass green or bluish green in KOH solution was discovered in northern California, USA. The new species is associated with Monterey cypress (Cupressus macrocarpa Hartw. ex Gord.) and redwood (Sequoia sempervirens (D. Don) Endl.), two tree species endemic to coastal California. The species is described
70
below and compared with phenetically similar taxa. Color terms and notations are from Kornerup and Wanscher (1978). Spore statistics include: x, the arithmetic mean of the spore length by spore width (+ SD) for n spores measured; Q, the quotient of spore length and spore width in any one spore, indicated as a range of variation in n spores measured; Q_, the mean of Q values.
Pseudobaeospora stevensii Desjardin sp. nov. Figures 1-7
Pileus 4-20(-26) mm latus, obtuse conicus vel campanulatus, siccus, glabrus vel subrugosus, brunneus vel obscure castaneus, 3% KOH ope azureo-viridis. Lamellae subliberae, angustae, brunneae vel pallide aurantiobrunneae. Stipes 20-35 X 1-3 mm, cylindricus, siccus, apicaliter ex furfuraceo scabrosus, basin versus squamulis brunneis vel aurantiogriseis instructus, basaliter albidulostrigosus. Colores violacei desunt. Basidiosporae 3.2-4 X 2.8-3.2 pm, ovoideae, subglobosae vel late ellipsoideae, leves, hyalinae vel pallide stramineae (in aqua), forte dextrinoideae, haud metachromaticae, membrana subincrassata instructae. Basidia 4 spora. Cheilocystidia 27-38 X 3.5-8 jum, cylindrica vel clavata, hyalina, tenuitunicata. Pleurocystidia nulla. Pileipellis ex hyphis cylindricis appressisque, 3-8(-10) um diam. cutem formantibus, haud_ gelatinosis, membrana dextrinoidea, hyalina vel brunnea (pigmento intraparietali vel incrustato) instructis. Hypodermium subcellulare, haud gelatinosum. Caulocystidia 40-80 X 4-7 jum, cylindrica vel clavata. Fibulae praesentes. Caro 3% KOH ope olivacea vel azureo-viridis. Ad frustula dejecta sub Cupresso et Sequoiae. USA (California). Holotypus hic designatus: A.W. Wilson #133 (SFSU).
Etymology: Named in honor of Fred Stevens who discovered the new species and who has been instrumental in documenting the fleshy fungi of California.
Pileus (Figs. 1-2) 4-20(-26) mm diam, obtusely conical to campanulate, sometimes broadly campanulate at maturity with uplifted margins; margin even to striate or short- sulcate in age; surface dry, glabrous, smooth to wrinkled; when young disc dark brown to dark reddish brown (8-10F5-7) with a slightly paler reddish brown margin (8-9DS5-7), in age becoming paler overall to light brown (7D5-6) or retaining a dark brown margin (5-8F8), lacking any violet or lavender tones; drying dark brown (7F5-8) overall; 3% KOH on pileus surface turning bluish green. Context 0.5-1 mm thick, pallid. Odor fungal. Taste slightly acrid and peppery. Lamellae subfree, close with 3-4 series of lamellulae, broad (3-4.5 mm), convex to ventricose, brownish orange (7C5) to brown (7D4-5), lacking violet or lavender tones; drying brown (7D-E5). Stipe 20-35 X 1- 3 mm, central, cylindrical, fistulose, dry, apex furfuraceous to scabrous, squamulose elsewhere, base covered with long, strigose, off-white to pale orangish white (5-6A-B2) hairs; apex brown (7E7-8), base dark brown (7F7-8), with greyish orange (6B3) to pale brownish grey (6C3) squamules.
4d
Fic. 1. Basidiomes of Pseudobaeospora stevensii (AWW 133-— Holotype). Scale: Twice natural
size. A color photograph of this specimen may be viewed at www.mykoweb.com.
Basidiospores (Fig. 3) 3.2-4 X 2.8-3.2 wm [x = 3.45 + 0.26 X 3.0 + 0.1 wm, Q = 1.1-1.3, Q. = 1.14 + 0.08, n = 25 spores), ovoid to subglobose or broadly ellipsoid with an eccentric hilar appendage, smooth, hyaline to pale yellow in H,0, strongly dextrinoid, non-metachromatic, with relatively thick walls. Basidia (Fig. 4) 16-20 X 4.5-5 ym, clavate, 4-spored, rarely 2-spored, clamped. Basidioles (Fig. 4) clavate. Cheilocystidia (Fig. 5) common, lamellar edge sterile or heteromorphous with clusters of cheilocystidia, 27-38 X 3.5-8 jm, cylindrical to narrowly clavate or irregular in outline, hyaline, thin-walled, greenish in 3% KOH. Pleurocystidia absent. Pileipellis (Fig. 6) two-layered: suprapellis a thin (up to 30 wm thick) cutis of appressed, radially arranged, cylindrical hyphae 3-8(-10) jm diam, with repent to erect, cylindrical to clavate or fusoid terminal cells; walls non-gelatinous, dextrinoid, hyaline to brown in H,O and 3% KOH; pigment intraparietal and incrusting; suprapellis overlaying a well- developed subpellis. Subpellis (Fig. 6) composed of inflated to irregular or puzzle-like hyphae (subcellular) 6-24 wm diam, forming a layer up to 100 wm thick; hyphae non- gelatinous, dextrinoid, hyaline to pale yellowish brown in H,O, olive to grass green or bluish green in 3% KOH, pigment intraparietal. Pileus trama of interwoven, elongated, cylindrical hyphae 2.5-6 7m diam, non-gelatinous, weakly dextrinoid, hyaline to yellow in H,O, vaguely green in 3% KOH.
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Fics. 2-7. Pseudobaeospora stevensii (AWW 133- Holotype). 2. Basidiomes. 3. Basidiospores. 4. Basidia and basidioles. 5. Cheilocystidia. 6. Pileipellis and hypodermium. 7. Caulocystidia. Scale bar: 2 = 10 mm; 3-5 = 10 wm; 6-7 = 20 ym.
fs
Hymenophoral trama regular; hyphae 3-9 wm diam, similar to those in pileus trama. Stipe tissues monomitic. Stipitipellis a cutis of repent hyphae with clusters of erect caulocystidia; cortical hyphae 2.5-4 wm diam, non-gelatinous, dextrinoid, hyaline to pale brown in H,O, olive to green in 3% KOH; medullary hyphae 2.5-10 pm diam, similar to cortical hyphae. Caulocystidia (Fig. 7) common, 40-80 X 4-7 pm, cylindrical to narrowly clavate, similar to the cheilocystidia, hyaline, thin-walled. Clamp connections present in all tissues.
Habit, habitat and distribution — Scattered to gregarious in leaf litter under Monterey cypress (Cupressus macrocarpa) or rarely under redwood (Sequoia sempervirens). Aug.-Dec. California.
SPECIMENS EXAMINED - UNITED STATES. CALIFORNIA: Marin
Co., Audubon Canyon Ranch, Galloway Canyon, 11 Dec. 1979, C. Calhoun 79-
1255. San Mateo Co., San Bruno, Skyline College campus, | Sept. 2001, coll.
by F. Stevens, AWW /33 (Holotype); same location, 26 Aug. 2001, Stevens 8-
26-01; same location, 10 Sept. 2001, Stevens 9-10-01. All specimens deposited
in. SHSU. Commentary — Pseudobaeospora stevensii is characterized by basidiomes that are dark brown overall (lacking any violet, lilac or lavender colors), and have very small, dextrinoid basidiospores, a two-layered pileipellis (with filamentous suprapellis and subcellular subpellis), 4-spored basidia, conspicuous narrowly clavate cheilocystidia and caulocystidia, and abundant clamp connections. It has been collected only in association with cypress and redwood in California. Within the genus Pseudobaeospora, the new species is unusual in forming dark brown basidiomes with tissues that turn olive to grass green or bluish green in 3% KOH. This microchemical reaction is reminiscent of that exhibited by a number of Gymnopus species allied with G. alkalivirens (Singer) Halling (cf. Halling 1979, 1981, 1990), and Leucoagaricus species (E. Vellinga, pers. comm.). Bas (2002, 2003) reports five other alkaline-virescent species of Pseudobaeospora; viz., P. dichroa Bas, P. pallidifolia Bas, A. Gennari & Robich, P. pyrifera Bas & L.G. Krieglst., P. jamonii Bas, Lalli & Lonati, and P. mutabilis Adamcik & Bas. The latter five species differ from P. stevensii by the following features. Pseudobaeospora dichroa differs in forming basidiomes with purplish brown to violet pilei and lamellae, lacks the two-layered pileipellis, and more importantly has a KOH reaction that is initially red, fading to yellowish green in time (Bas 2002, 2003). Pseudobaeospora pallidifolia differs in forming violet brown pilei with white to pinkish cream lamellae, larger basidiospores (4.5-6 X 3-4.5 um), lacks cheilocystidia and lacks the two-layered pileipellis (Bas et al. 1997; Bas 2003). Pseudobaeospora pyrifera differs in forming dark purplish to vinaceous brown or pinkish brown pilei, reddish violaceous lamellae, a dark reddish brown stipe, has broader cheilocystidia (up to 13 wm diam.), and also lacks the two- layered pileipellis (Bas and KriegIsteiner 1998; Bas 2003). Pseudobaeospora jamonii, shares with P. stevensii the two-layered pileipellis, narrowly clavate cheilocystidia and
74
small basidiospores, but differs significantly in forming greyish purple to violaceous brown pilei, lilac to yellowish grey lamellae, a lilac to purplish brown stipe, has a paler green KOH reaction, and is associated with Alnus, Fraxinus, Fagus, Corylus and Picea (Bas et al. 2002; Bas 2003). Pseudobaeospora jamonii is known at present only from northwestern and central Italy. Pseudobaeospora mutabilis differs in forming dark violaceous grey-brown pilei with pinkish grey margins, violet lamellae, broadly clavate cheilocystidia 7-11(-18) wm in diam., and a hymeniform pileipellis of erect chains of inflated cells. The latter species is associated with Molinia, Frangula and Betula and is known at present only from the type locality in Slovakia (Adamcifk and Bas 2002; Bas 2003).
Notes on North American Pseudobaeospora — Few species of Pseudobaeospora have been reported from temperate North America. Coker (1929) reported Prunulus syringeus Murrill from North Carolina. It was described originally from Jamaica, and is now known as Pseudobaeospora murrillii E. Horak (1964; a new name, non Pseudobaeospora syringea Singer). Smith (1947) mentioned this species (as Mycena syringea (Murrill) Murrill) and suggested that it was similar to Tricholoma microsporum Ellis, which he collected in Michigan, New York, Ontario and California. Although Smith studied the type specimen of Pr. syringeus (NY), the holotype packet is now empty so I was unable to obtain any microscopic data (pers. obs.). I have collected numerous specimens of what Coker (1929) called Pr. syringeus from North Carolina, Tennessee and other areas in the Appalachian Mts., and the latter species forms deep violet basidiomes with abundant clamp connections, no cheilocystidia, has a cutis-type pileipellis of hyphae 5-12 wm diam, lacks a subcellular hypodermium, and does not discolor in KOH. Whether this is the same species as that from Jamaica cannot be determined at present. The type of Tricholoma microsporum was distributed as part of the Ellis and Everhart, North American Fungi Exsiccata, as #2003, from Newfield, New Jersey. I have studied several representative specimens (K, NY, SFSU) and this species forms violet basidiomes with a thin cutis-type pileipellis that overlays a subcellular hypodermium, has dextrinoid basidiospores 4-5 X 3.5-4.5 yum, lacks cheilocystidia, lacks clamp connections, and does not discolor in KOH. Clearly, it is distinct from Pr. syringeus sensu Coker.
Agaricus fuscolilacinus Peck (1886) was described from the Adirondack Mts. of New York, and was reported to form watery-brown to lilac-brown, glabrous pilei, close, adnexed, brownish lamellae, and a brown stipe with pruinose apex and white, villose hairs at the base. An analysis of the holotype specimen (NYS!) revealed the following features: Basidiospores 4.2-6.8 X 3.2-4 wm [x = 5.2 +0.6 X 3.5 +0.2 wm, Q= 1.2-2,Q. = 1.5+0.2, n = 30], ellipsoid to lacrymoid, thick-walled, smooth, inamyloid to weakly dextrinoid. Basidia 16-20 X 4-5 um, 2-spored. Cheilocystidia absent. Pileipellis a hymeniform layer of broadly clavate cells 12-26 X 7-14 wm, sometimes in chains, thin- walled, subhyaline to pale ochraceous in KOH, inamyloid. Tramal hyphae 4-12 wm
75s)
diam, hyaline, inamyloid, thin-walled, non-gelatinous. Stipe vesture a tangled layer of hyaline to pale yellow hyphae 2-5 wm diam, cylindrical, inamyloid, with a few erect, cylindrical to vesiculose caulocystidia. Clamp connections present. This species belongs in Pseudobaeospora where it is allied with P. mutabilis. A formal transfer will not be made until more specimens from the Adirondack Mts. referable to A. fuscolilacinus are collected for comparison. The latter species differs from P. stevensii in forming larger basidiospores on 2-spored basidia, in lacking cheilocystidia, and in lacking a two-layered pileipellis.
Pseudobaeospora pillodii (Quél.) Wasser was reported from British Columbia, Canada by Redhead (1982). Bas (2003) suggested that Redhead’s material may represent P. oligophylla (Singer) Singer, a species accepted by Redhead as a synonym of P. pillodii. These two taxa differ only subtly in pileipellis anatomy. Nonetheless, both taxa are quite distinct from P. stevensii in forming lilac to purple basidiomes that lack cheilocystidia, lack clamp connections, and do not discolor to green in KOH.
Acknowledgments
I would like to thank Dr. Fred Stevens for bringing this new species to my attention and for all of his fine contributions to mycology in California. I thank Andrew Wilson for taking notes on fresh material of the holotype specimen, Dr. Egon Horak for correcting the Latin diagnosis, and Dr. Else Vellinga and Brian Perry for reviewing the manuscript and for providing helpful comments and obscure literature.
Literature Cited
Adamcik S, Bas C. 2002. Pseudobaeospora mutabilis, a new species discovered in Slovakia. Mycotaxon 84: 272-275.
Bas C. 2002. A reconnaissance of the genus Pseudobaeospora in Europe I. Persoonia 18: 115-122.
Bas C. 2003. A reconnaissance of the genus Pseudobaeospora in Europe II. Persoonia 18: 163-199.
Bas C, Gennari A, Robich G. 1997. Pseudobaeospora pallidifolia Bas, Gennari & Robich. Una nuova specie dall’Italia. Riv. Micol. 40(3): 195-199.
Bas C, Krieglsteiner L. 1998. Pseudobaeospora pyrifera, a new species found in southern Germany and The Netherlands. Z. Mykol. 64(2): 203-206.
Bas C, Lalli L, Lonati G. 2002. Pseudobaeospora jamonii, una nuova specie in Italia. Micol. Veget. Medit. 17(1): 31-35.
Coker WC. 1929. Notes on fungi. J. Elisha Mitchell Sci. Soc. 45: 164-178.
Halling R. 1979. Notes on Collybia. I. Collybia alkalivirens. Mycotaxon 8: 453-458.
Halling R. 1981. Notes on Collybia. I. Additional taxa that are green in alkaline solution. Mycologia 73: 634-642.
Halling R. 1990. Collybia fuscopurpurea in the Americas. Mycol. Res. 94: 671-674.
Horak E. 1964. Notes sur Pseudobaeospora Singer. (1942), excl. Lepiota, sect. Sericellae. Rev. Mycol. France 29: 72-81.
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Kornerup A, Wanscher JH. 1978. Methuen handbook of colour. 3rd Ed. Eyre Methuen, London. 252 p.
Peck CH. 1886. Report of the botanist. Ann. Report N.Y. State Mus. 39: 30-73.
Redhead SA. 1982. Pseudobaeospora pillodii. Fungi Canadensis 217.
Singer R. 1938. Notes sur quelques Basidiomycétes. Rev. Mycol. France 3: 187-199.
Singer R. 1942. Type studies on agarics. Lloydia 5: 97-135.
Singer R. 1963. The delimitation of the genus Pseudobaeospora. Mycologia 55: 13- ie
Smith AH. 1947. North American species of Mycena. Univ. of Michigan Press, Ann Arbor, Michigan. 521 p.
MYCOTAXON
Volume 90(1), pp. 77-80 July-September 2004
Les pycnides et conidies de Lecanora vaenskaei
(lichens, Lecanoraceae)
CLAUDE Roux
claude.roux2 1 @wanadoo.fr CNRS, UPRES A 6116, Laboratoire de botanique et écologie méditerranéenne, Institut méditerranéen d’écologie et de paléoécologie, Faculté des sciences et techniques de Saint-Jéréme, FR—13397 MARSEILLE cedex 20, France.
Résumé—La découverte de pycnides sur un spécimen de Lecanora vaenskaei permet de compléter la description de cette espéce et de confirmer sa position systématique. Elle est comparée avec les espéces voisines, plus particuli¢érement L. achariana.
Abstract— Discovery of pycnidia on a specimen of Lecanora vaenskaei confirms its systematic position. The species description is emended, and the species is compared to related species, especially L. achariana.
Resumo—La malkovro de piknidioj en specimeno de Lecanora vaenskaei ebligas kompletigi la priskribon de tiu—chi specio kaj konfirmi ties sistematikan lokon. Komparo kun la parencaj specioj, pli speciale L. achariana.
Introduction
Lecanora vaenskaei Cl. Roux et Clother Coste, décrit par Roux et al. (1993), est un Lecanora saxicole-calcifuge du sud-est méditerranéen francais caractérisé par un thalle épilithique vert-jaune clair, formé de granules se transformant en petites squamules ombiliquées courtement lobées, a cortex supérieur et cortex inférieur, et contenant de l’acide usnique et de la zéorine. Les apothécies qui se forment a |’extrémité des lobes des squamules sont typiquement lécanorines et nettement resserrées a la base ou méme courtement pédonculées. Par la structure de ses apothécies, Lecanora vaenskaei appartient bien aux Lecanoraceae, tout comme des espéces voisines (L. chaffiniana Houmeau et Cl. Roux, L. chlorophaeodes Nyl., L. weberi Ryan) et les Rhizoplaca (Roux et al. 1993).
Quoique trés détaillée et richement illustrée, la description originale de L. vaenskaei est incompléte, puisque ses auteurs n’avaient pas observé de pycnides. Or, en réexaminant un spécimen (Roux n° 21429), j’ai eu la chance d’y découvrir quelques pycnides typiques, riches en conidies, dont |’ observation microscopique a permis de compléter la description de |’espéce et de confirmer Sa position systématique. Ce spécimen est choisi comme épitype.
78
Methodes Les coupes et préparations microscopiques ont été réalisées 4 main levée et observées dans l’eau et le bleu coton au lactophénol, avec un microscope photonique (grandissement maximal de x 1500) muni d’un dispositif de contraste interférentiel. Les dessins ont été réalisés avec l’aide d’un tube a dessin. Le seul spécimen disponible étant ancien, les mesures ont été effectuées dans |’eau, sur du matériel mort.
Lecanora vaenskaei Cl. Roux et Clother Coste
in Roux, Coste, Ménard, Bellemére et Bricaud, Can. J. Bot., 71: 1660-1671 (1993). France, Provence, Var, Le Muy, la Roquette, 500 ma |’ouest de la chapelle ruinée, sur une surface rocheuse de bréche non calcaire, périodiquement inondée. Alt. 110 m. 1992/10/29. Holotype MARSSJ; isotypes MARSSJ, GZU, H.
Epitype: France, Provence, Var, Roquebrune-sur-Argens, rocher de Roquebrune, partie E, prés du chemin, sur une surface rocheuse de bréche non calcaire. Alt. 60 m. 1992/05/21. MARSSJ, Herbier C. Roux n°21429.
Pycnides compléetement enfoncées dans le thalle (situées dans la couche algale), enti¢rement incolores, subglobuleuses, d’environ 0,25-0,3 x 0,2- 0,25 mm, simples (uniloculaires). Les coupes ne passant pas par l’ostiole celui—ci n’a pu étre observé, mais il n’est pas saillant puisque non visible au stéréomicroscope. La pycnide (fig. 1) est délimitée par une mince paroi incolore de 5—10 wm d’épaisseur, formée de deux couches de cellules globu- leuses ou subglobuleuses de 3,5-5,5 x 2,5-4,5 wm de diameétre (lumiére de 1-2,5 x 1-2 ym), a paroi épaisse (de 1-2 ym). Conidiophores simples ou ramifiés, a cellules de 3-10,5 x 2—3 wm (lumiére de 2—9,5 x 1-2 ym, paroi d’environ 0,5 wm d’épaisseur), de type exobasidié (portant la ou les cellules conidiogénes 4 leur ou leurs extrémités distales). Cellules conidiogénes (fig. 1 et 2) longuement ampulliformes, de 6,5—11 x 2,5-—3,5 wm (lumiére de 5— 9,5 x 1-2 ym, paroi de 0,5—-1 wm d’épaisseur). Conidies (fig. 3) filiformes, modérément courbes, de (16)18—31 x 1 ym (d’aprés 16 mesures).
Discussion
L’étude de la morphologie et de la structure des pycnides et des conidies de L. vaenskaei confirme la parenté de cette espéce avec les Lecanora a thalle squamuleux ou placodiomorphe jaune verdatre a acide usnique, en particulier L. achariana A. L. Sm., L. chaffiniana, L. chlorophaeodes et L. dispersoareolata (Schaer.) Lamy. Dans ce groupe d’espéces, L. chlorophaeodes et L. dispersoareolata s’éloignent le plus de L. vaenskaei par |’absence de cortex inférieur méme rudimentaire, tandis que L. chaffiniana a une ébauche de cortex (Houmeau et Roux 1988) pigmentée de vert noiratre et L. achariana, tout comme L. vaenskaei, un cortex inférieur non pigmenté ou pigmenté de brun rouge ou de rose au contact du support, cortex déja observé par Harmand
Y
(1913: 926). L. chaffiniana se distingue également de L. achariana et de L. vaenskaei par la présence d’acide gyrophorique et |’absence de zéorine.
Bien que Roux et al. (1993) n’aient pas comparé L. vaenskaei avec L. achariana, c’est pourtant cette espéce qui a le plus d’affinités avec L. vaenskaei, notamment par la structure du thalle, la morphologie et la structure des apothécies, par des conidies (Harmand 1913: 927) d’environ 30 x 1 ym et par son écologie (sur des surfaces d’écoulements temporaires). L. vaenskaei s’en distingue néanmoins (tableau 1) par un thalle nettement plus adhérent au substrat, d’aspect plus granuleux que squamuleux, a lobes beaucoup plus petits, des apothécies terminales, des spores plus larges et l’absence de triterpénoides.
10 um
10 um
Fig. 1. Structure microscopique de la partie basale d’une pycnide de Lecanora vaenskaei d’apres une coupe radiale observée dans le bleu coton au lactophénol. a: cellule algale; c: conidie; ca: couche algale (hyphes et cellules algales); cc: cellule conidiogéne ; cp: conidiophore; p: paroi de la pycnide formée de deux couches de cellules a paroi €paisse.
Fig. 2. Cellules conidiogénes de Lecanora vaenskaei, portant une conidie immature, observees dans le bleu coton au lactophénol. a: cellule conidiogéne; c: conidie.
Fig. 3. Conidies matures de Lecanora vaenskaei, observées dans |’eau.
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Tableau 1. Caractéres distinctifs de Lecanora vaenskaei et L. achariana
oie eiasbaaadon Lecanora vaenskaei Lecanora achariana
nettement adhérent au support, d’aspect plus granuleux que squamuleux, formé
Thalle de granules ou de petites squamules a lobes de 0,3—0,5 x 0,15-0,3 mm
peu adhérent au support, d’ aspect nettement squamuleux, formé de squamules a lobes de 1,5 x 0,5-2 mm
(9)10-14(17) x 10-15(16) x
seal 4(4)6-8(9)10 pm 4-6 wm
acide, usnique, zéorine, triterpénoides (Purvis et al. 1992)
acide usnique, zéorine (Roux Chimisme et al. 1993)
montagnes d’ Europe centrale et septentrionale, de Corse
et de Sardaigne; étage montagnard; non thermophile
région méditerranéenne; étage mésomméditerranéen; thermophile
Répartition et écologie
Spécimen de Lecanora achariana examiné: France, Haute-Corse, forét d’Aitone, prés du foyer de ski de fond, sur une surface subhorizontale de granite tres cohérent, soumise a de brefs écoulements périodiques. Alt. 1100 m. 1999/04/19. MARSSJ, Herbier C. Roux (sans numéro).
Bibliographie
Harmand J, 1913. Lichens de France. Catalogue systématique et descriptif. Crustacés. Pannariés, heppiés, lécanorés, pertusariés, thélotremés. L. Lhomme édit., Paris, p. 761-1185.
Houmeau JM, Roux C, 1988. Lecanora chaffiniana Houmeau et Roux sp. nov., espéce nouvelle de lichen du Massif central (France). Bull. Soc. bot. Centre-Ouest, nouv. sér., 19: 265-277.
Purvis OW, Coppins BJ, Hawksworth DL, James PW, Moore DM, 1992. The lichen flora of Great Britain and Ireland. Natural History Museum Publications et British Lichen Society édit., London, 710 p.
Roux C, Coste C, Ménard T, Bellemére A, Bricaud O, 1993. Lecanora vaenskaei Roux et Coste sp. nov. (Lichenes, Ascomycotina), sa position systématique et celle des Rhizoplaca (Lecanorales, Lecanoraceae). Can. J. Bot., 71: 1660-1671.
MYCOTAXON
Volume 90(1), pp. 81-123 July-September 2004
New and previously unrecorded saxicolous species of Buellia s.|. with one-septate ascospores from the Greater Sonoran Desert Region
FRANK BUNGARTZ
bungartz@bsm.mwn.de Botanische Staatssammlung Miinchen Menzinger Strafpe 67 D-80638 Miinchen, Germany
Abstract—The following species of Buellia s.1. from the Greater Sonoran Desert Region are described new to science: Buellia nashii, B. navajoensis, B. regineae and B. sheardii. A valid description is provided for B. lepidastroidea, a name introduced by Imshaug, but never validly published. Buellia argillicola 1s reported from the Sonoran Region for the first time. A single record of the rare and unusual B. vilis is reported from Arizona. Buellia novomexicana and B. fusca are synonymized with B. tyrolensis. Detailed descriptions are provided for all these species. In addition a dichotomous key includes all saxicolous species with one-septate ascospores currently known from the Greater Sonoran Desert Region.
Zusammenfassung-Die folgenden neuen Arten der Sammelgattung Buellia s.1. werden aus der Sonoraregion beschrieben: Buellia nashii, B. navajoensis, B. regineae and B. sheardii. Eine weitere Art ist Buellia lepidastroidea. Diese Art wurde bereits von Imshaug vorgeschlagen, aber nicht giiltig publiziert. Sie wird hier erstmals mit einer giiltigen Artdiagnose versehen. Buellia argillicola wird erstmals aus der Sonoraregion nachgewiesen. Ein Einzelfund der seltenen und ungewohnlichen Art B. vilis wird aus Arizona gemeldet. Buellia novomexicana und B. fusca werden mit B. tyrolensis synonymisiert. Alle diese Arten sind hier detailliert beschrieben. Dariiber hinaus erm6glicht ein dichotomer Schliissel die Bestimmung aller bisher aus der Region bekannten, saxikolen Arten mit zweizelligen Sporen.
Key Words—taxonomy, lichenized ascomycetes, Physciaceae, new _ species, dichotomous key
Introduction
A revision of the saxicolous species of Buellia s.1. with one-septate ascospores from the Greater Sonoran Desert Region currently accepts a total of thirty-one species. Several of these species have never been reported from the area or are not yet described. Descriptions of these new species and records are given here. A complete treatment of the genus will be included in the third volume of the Sonoran Desert Lichen Flora (Nash et al. 2002) but a key to all species is published here to allow identification of all species currently known from this region.
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The species described here are not necessarily members of what might be considered the core group of Buellia De Not. (Bungartz & Nash 2004c). Traditionally specimens, which are related to Buellia aethalea (Ach.) Th. Fr. have been associated with Buellia s.str. but the current Botanical Code (Greuter et al. 2000) lists Buellia disciformis (Fr.) Mudd, a species with distinct characters of Hafellia Kalb, H. Mayrhofer & Scheid., as the type. There is currently some discussion to change this typification, but a proposal by Moberg et al. (1999) is not presently accepted. Until the outcome of this proposal is decided, a broad concept of Buellia s.1. is therefore adopted. Taxonomic affinities of the species described, are discussed in more detail within brief notes for each species. The genus concept of Buellia is currently in a state of much disarray (Eriksson et al. 2002, Nordin 2000) and the question, which genera will ultimately be recognized should best be addressed using molecular tools to evaluate classical data.
Methods
All specimens were examined with light microscopy using hand- and cryosections. Both conventional bright field microscopy (BF) as well as differential interference contrast (DIC) were used. Selected specimens were also studied with transmission electron microscopy (TEM) according to a protocol described in detail by Bungartz et al. (2002). To improve dehydration and infiltration this protocol has been modified according to Bungartz & Nash (2004a).
All specimens were spot tested and routinely examined with standardized thin-layer chromatography (Culberson & Johnson 1982, Culberson & Kristinsson 1970, Orange et al. 2001, White & James 1985). TLC-Plates were interpreted with the computer program WINTABOLITES (Mietzsch et al. 1994), and scanned for permanent record Egan (2001). In addition a subset of specimens was analyzed by Dr. Jack A. Elix from the Australian National University in Canberra using standardized High Performance Liquid Chromatography (HPLC, Feige et al. 1993). Spores measurements are given according to Nordin (2000). Pigment names follow Meyer & Printzen (2000). Detailed specimen information about all collections deposited at ASU is available at: http//seinet.
asu.edu/collections/selection. jsp.
Taxonomic Descriptions
Buellia argillicola de Lesd., Ann. Cryptog. Exot. I: 243 (1929) Figure 1
Type: MEXICO. Mexico. San Jerénimo. On volcanic rocks (pebbles embedded in soil) [original label data: Estado de Distrito Federal. San Jer6nimo, Sobre Piedra], 19°13’30”N, 100°01°30”W, 2598 m alt., Pedro (Pierre) Lyonett no. 110 (MEXU! — neotype selected
here).
Taxonomic note: No type material from the original collections was found. It can be assumed that de Lesdain’s holotype collection, originally located in Dunkerque (France), has been
destroyed during the Second World War. The specimen selected as a neotype was collected at
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the type locality and agrees well with the description in the protologue.
THALLUS (Fig. 1A, B)—Crustose, thin, usually growing in distinct circular patches, subeffigurate, several thalli often confluent, epilithic, continuous to rimose with fine fissures; prothallus distinct, delimiting the thallus as a black outline; thallus surface matt and chalky, dull, usually white, rarely gray, heavily pruinose, phenocorticate; entire thallus filled with an abundance of calcium oxalate (H,SO,+ forming clusters of
needle shaped crystals).
APOTHECIA—Lecideine, (0.3—)0.5—0.8(—1.1) mm in diameter, remaining immersed
Fig. 1: Light micrographs of Buellia argillicola (Nash 12152).— A. Overview of the rimose
thallus. —B. Close-up of the rimose thallus and apothecia.—C. Cross section of the aethalea-
type exciple.—D. Oblong, ascospores. —E. Ascospore: no septum thickening is present during
the spore ontogeny.
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to indistinctly adnate; proper margin thick, persistent, not excluded with age, black, usually covered with a dense, fine, white pruina; disc black, plane, not becoming convex with age, usually covered with a dense, fine, white pruina; proper exciple similar to the aethalea-type (Fig. 1C) sensu Scheidegger (1993), i.e. inner exciple almost entirely reduced, hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), similar in structure and orientation to the paraphyses, transient with the deep reddish brown hypothecium (leptoclinoides-brown, textura intricata), outer exciple expanded, globular cells strongly swollen (+ textura oblita) and moderately carbonized with a brown pigment (cf. elachista-brown), pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown (cf. elachista-brown) pigment cap.
ASCI—8-spored, clavate, Bacidia-type. ASCOSPORES (Fig. 1D, E)—oblong to ellipsoid, usually not constricted, with obtuse ends, not curved, (12.0—)15.0-[16.8]- 18.5(—20.0) x (6.0—)6.5—[7.3]-8.1(—9.0) wm (n= 60), one-septate, proper septum narrow, not thickening during spore ontogeny, lateral wall thickenings absent [Beltraminea
(=Buellia)-type]; ornamentation rugulate.
PYCNIDIA—Rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980 and Vobis & Hawksworth 1981); conidia simple, bacilliform, 4.0—6.5 x 1.0-1.5 wm (n = 20).
CHEMISTRY—With the depsidones norstictic, connorstictic, stictic and hypostictic acid and one unknown substance (RF 7, not visible in natural light, UV+ orange). Thallus and medulla K+ faint yellow (yellow solution, but not orange, needle-shaped crystals observed in the microscope), P+ faint yellow, C-, KC—, CK—. UV+ pale white. Thallus and apothecia react amyloid with Lugol’s (always test with concentrated Lugol’s iodine or in the compound microscope; positive reactions can be very weak!).
SUBSTRATE AND ECOLOGY—The type specimen grows on pebbles embedded in calcareous (HCI+) soil (= tépetate), the only known Sonoran specimens grow on limestone (strongly HCI+).
DISTRIBUTION (Fig. 3)—Rare, in the Sonoran Desert only known from a single location, originally described from central Mexico.
NOTES—Specimens of B. argillicola are superficially similar to B. subalbula (Nyl.) Miill. Arg. or B. venusta (K6rb.) Lettau. The thalli of B. argillicola are, however, more continuous and less extensively fissured. Apothecia of B. argillicola have a plane disc with a persistent lecideine margin and no thalline collar or veil. Internally the structure resembles the aethalea-type but the outer exciple is much more prominent and persistent. Both B. subalbula and B. venusta have a thin, largely reduced exciple, which
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usually becomes excluded by the swelling of the disc. In B. venusta a thalline collar is common and it can sometimes be present in B. subalbula. Buellia subalbula is restricted to the coast whereas B. argillicola is known from inland localities. Buellia subalbula is characterized by a fuscous brown to deeply aeruginose exciple, which always reacts HNO,+ violet. Buellia venusta and B. argillicola have no aeruginose exciple pigment. Buellia venusta has pluriseptate, B. argillicola one-septate ascospores.
REPRESENTATIVE SPECIMEN EXAMINED—MEXICO. Sinatoa. Nash /2/52 (ASU).
Buellia lepidastroidea Imshaug ex Bungartz sp. nov. FIGURE 2
Thallus saxicolus, crustaceus, areolatus vel bullato-areolatus, crassus, eburneus, cum hypothallo atero. Apothecia sessilia, lecideina, marginibus propriis exclusis. Excipulum tenuis, fulvum, sine pigmento aeruginoso, carbonaceum. Hymenium interspersum guttae olei. Asci 8-spori. Sporae uniseptae, oblongae vel ellipsoideae, septibus incrassatus, 7-17 x 5-8 pm. Pycnidia urceolata vel globosa. Conidia bacilliformes, 2-7 x 1-2 ym. Thallus atranorina,
+ diploicina et + fulgidina continens. Medulla non- amyloideus.
Type: MEXICO. Basa Ca.irorniA. Isla Cedros, ridge crest overlooking western shore and adjacent canyon to the E at the NW corner of the island, 28°22’00”N, 115°15’30”W, ca. 300 m alt., on acidic rock, 19 March 1994, Nash 34458 (ASU! — holotype designated here).
Taxonomic note: The species name was first used by Imshaug (1951) in his dissertation but
was never validly published.
THALLUS (Fig. 2A, B)—Crustose, thin to moderately thickened, + continuous or becoming dispersed, epilithic; areolate to subsquamulose or bullate; prothallus delimiting the thallus margin, distinctly blackened to pale gray, rarely white and indistinct; thallus surface matt and dull, not shiny, usually ivory, rarely beige, epruinose or rarely with a fine pruina, phenocorticate; without Ca-oxalate crystals in the medulla (H,SO,-).
APOTHECIA—Lecideine, (0.3—)0.5—0.8(—1.1) mm in diameter, soon sessile; proper margin thin, black, rarely persistent, usually excluded with age; disc black, epruinose, plane, soon becoming convex with age; proper exciple of the dispersa-type (Fig. 2C) sensu Scheidegger (1993), i.e. inner excipular hyphae distinct, not reduced, pigmented, prosoplectenchymatous (fextura oblita), extending from the deep reddish brown hypothecium (leptoclinoides-brown, textura intricata); outer excipular hyphae short- celled, cells angular, distinctly swollen (textura angularis) and usually + carbonized with various amounts of brown pigments (cf. elachista-brown), pigmentation continuous with the epihymenium; hymenium hyaline, inspersed with oil droplets; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown).
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ASCI—$%-spored, clavate, Bacidia-type (Fig. 2F). ASCOSPORES (Fig. 2D, G, H)—oblong to ellipsoid, usually not constricted, with obtuse ends, not curved, (7.0—) 11.4—[13.2]-15.0(-17.0) x (5.0-)6.0-[6.7]-7.4(-8.0) um (n = 60); one-septate, proper septum briefly thickened but soon becoming reduced during spore ontogeny, lateral wall thickenings absent (+ Physconia-type); ornamentation absent (not visible in DIC); septum with septal pore canal, simple pore and undifferentiated pore plug; spore wall (Fig. 2G, H) differentiated into smooth, thin perispore (0.06—-0.07 um), narrow intermediate layer (< 0.03 wm), thick proper spore wall (0.34-0.54 wm) and moderately thickened endospore (0.12—0.21 ym).
PYCNIDIA—Rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980 and Vobis & Hawksworth 1981); conidia simple, bacilliform, 2.0—7.0 x 1.0—2.0 wm (n = 41).
CHEMISTRY—With the depsides atranorin and chloroatranorin and the depsidones diploicin, 3-dechlorodiploicin, fulgidin and isofulgidin. HPLC—artefacts: haematommic acid, methyl B-orsellinate (HPLC by J. Elix, Canberra). Thallus and medulla K+ yellow, P+ orange, C—, KC—, CK—. UV+ pale ivory. The thallus medulla and cortex does not react amyloid, apothecia are amyloid in Lugol’s (always test with concentrated Lugol’s iodine or in the compound microscope; positive reactions can be very weak!).
SUBSTRATE AND ECOLOGY—0n a variety of siliceous (HCI-) rock substrates.
DISTRIBUTION (Fig. 3)—Common along the coast of southern California and Baja California.
NOTES—tThalli of B. lepidastroidea sometimes vaguely resemble B. dispersa or
—>
Fig. 2: Buellia lepidastroidea (A-E. light micrographs; F-H. TEM micrographs).— A. Overview of the areolate, sublobate thallus with sessile apothecia (Nash 6742a).—B. Close-up of thallus and apothecia (Nash 6742a).—C. Cross section through the apothecium with inspersed hymenium (Nash 34458).—D. Premature ascospore with distinct septal thickening (Nash 34458).—E. Conidia (Nash 34458).—F. Bacidia-type ascus (Nash 38438, for designation of the different layers see Bellemére 1994): The a- and b-layer
(ab) are barely visible and cannot reliably be distinguished (possibly as a result of fixation artifacts); (c) outer electron opaque c-layer; (d1) d1-layer, i.e. the outer tholus, which is distinctly laminated (in light microscopy this outer part stains deep blue with Lugol’s iodine); (d2) d2-layer, i.e. the inner tholus, which is not layered and + homogeneous (not staining in Lugol’s iodine); (oc) ocular chamber .—G. Mature ascospore with reduced septal thickenings (Nash 38438).—H. Spore wall of a mature ascospore (Nash 38438): (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2) intermediate layer; (3) proper spore wall;
(4) endospore.
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B. nashii. Buellia lepidastroidea is, however, strictly coastal and characterized by oil droplets in the hymenium. This inspersion is usually distinct, especially if thick apothecial sections are treated with 5% KOH. In thin sections, oil inspersion may be less conspicuous but nevertheless always present. Thalli growing close to the seashore and subject to spray often show more exuberant growth than sheltered thalli. Their thallus areoles become strongly inflated and apothecia thus appear “stalked”.
The North American specimens fit well into the concept of Imshaug’s B. lepidastroidea and the species is described here to validate the name Imshaug first introduced for this species. The species belongs to a group of taxa with indistinct Physconia-type ascospores, a dispersa-type exciple and an oil-inspersed hymenium. This group of species is currently not well understood. Buellia lepidastroidea is very similar to European material of B. excelsa (Leight.) A.L. Smith. Scheidegger (1993) selected a lectotype for B. excelsa but the material could not be found at BM. Specimens identified as B. excelsa from Scheidegger’s private herbarium have a more distinctly brown thallus, contain only atranorin, not diploicin or fulgidin and were collected at high altitude, not close to the seashore. Scheidegger (1993, p. 349) mentions that the species is distributed “in dunes and in mountains”. In the Sonoran Desert Region B. lepidastroidea is clearly confined to coastal, maritime habitats and has not been found further inland or at high altitudes. Other taxa in the Buellia excelsa-group are B. boergesenii Imshaug and B. jorgensis Zahlbr. (Chile — M-0061335, Spindler 16/58, det. Follmann). Buellia conspirans (Nyl.) Vain. is a saxicolous species similar to B. boergensii according to Imshaug (1955). It is currently listed in the North American Checklist (Esslinger 1997) as a synonym of B. curatellae Malme. Buellia curatellae is, however, a corticolous species with distinct lateral spore wall thickenings. It was transferred by Marbach (2000) into the genus Hafellia.
Buellia lepidastroidea is treated here as a distinct species, but a more detailed revision of the Buellia excelsa-group is necessary to evaluate the taxa currently included in this group. Because of the oil inspersion in the hymenium, species of the Buellia excelsa— group show some affinities to the genus “Hafellia”. The presence of diploicin and related substances in B. lepidastroidea is unusual for Buellia s.str. but these substances have been reported from “Hafellia’. Scheidegger (1993) did, however, not include B. excelsa within “Hafellia” because the ascospores lack lateral wall thickenings and he did not find diploicin in B. excelsa.
Buellia lepidastroidea usually can be easily distinguished from B. (“Hafellia”’) regineae because of its thinner thallus, with smaller, more dispersed areoles, smaller apothecia and ascospores. Lateral wall thickenings are usually present in spores of B. regineae,
—> Fig. 3: Distribution of Buellia argillicola and Buellia lepidastroidea in the Sonoran Desert
(Floristic Provinces according to Shreve & Wiggins 1964).
Buellia lepidastroidea ‘
108 19° W
Subdivisions of the Sonoran Desert Floristic Province (Shreve & Wiggins 1964):
Arizona Upland
HL
Lower Colorado Region |
Magdalena Region
Plains of Sonora (including Foothils of Sonora)
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but less distinct than in B. leptoclinoides (see notes on B. regineae).
Thalli of B. lepidastroidea and B. excelsa are very similar to B. dispersa and B. nashii. All these species also share the same exciple anatomy and have indistinct Physconia- type ascospores. The angular thickening of the spore septum, which is characteristic for the Physconia-type ascospore, can only very briefly be observed in premature ascospores (Fig. 2D) but becomes soon reduced in mature spores (Fig. 2G & H). Buellia dispersa and B. nashii, however, do not contain oil droplets within the hymenium.
REPRESENTATIVE SPECIMENS EXAMINED—MEXICO. Baja Ca.irornia. Egan 13790 (OMA); Kalb 24657 (hb. Kalb); Scheidegger Inv. Nr. 170-189, 71-95, 150-155 (hb. Scheidegger); Weber L-43033, L-43014, L-43062, L-36575 (COLO); Wetmore 63852, 75765, 72431, 63498, 75873 (MIN); Nash 38242, 34370, 38241, 8688b, 34319, 4916, 34052, 38256, 8730, 25188, 38459, 34358, Ryan 2136, 21263 (ASU). Baya CALIFORNIA Sur. Wetmore 72223 (MIN); Marsh 6080, Nash 8947 (ASU).—UNITED STATES. Ca.irornia. Los Angeles Co. Bratt 9799, Bratt 10219, 11843, 11844 (SBBG); Hasse Exs. 155 (ASU); Weber L-42606, L- 42823 (COLO). Santa Barbara Co. Bratt 7379, 7605 (SBBG). Nash 32646, 33021, Ryan 31255b, 31270 (ASU). WasHINGTON. Kittitas Co. Ryan 16838 (ASU).
Buellia nashii Bungartz sp. nov. FIGURE 4
Similis Buelliae dispersae sed apotheciae cum pigmento aeruginoso et thallus acida norstictica
et connorstictica vel acida stictica continens. Etymology: The species is named in honor of my Ph.D. supervisor Dr. Thomas H. Nash III.
Type: MEXICO. CoanuiLa. 3 km W of the paved road, at the Delores sign, 26°00’00”N, 101°00’00”’W (estimated coordinates), on volcanic rock, 20 March 1976, Nash 6742a (ASU! — holotype designated here; one isotype at MEXU, one isotype at US).
THALLUS (Fig. 4A, B)—Crustose, thick, + continuous or becoming dispersed, epilithic; areolate to subsquamulose; prothallus absent; thallus surface matt and smooth to deeply fissured, dull or + shiny, usually ivory, beige to deep brown or gray, rarely lead gray, with fine or coarse pruina, rarely epruinose, phenocorticate; with few, or rarely large amounts of Ca oxalate crystals (H,SO,+ forming clusters of needle shaped sulphate crystals).
APOTHECIA—Lecideine, (0.4-)0.6—1.1(-1.2) mm in diameter, sessile; proper margin black, thin to thick, usually persistent, rarely excluded with age; disc black, usually epruinose, rarely with white pruina, plane, often becoming strongly convex with age; proper exciple of the dispersa-type (Fig. 4C) sensu Scheidegger (1993), i.e. inner exciple distinct, not reduced, hyphae pigmented, prosoplectenchymatous (textura oblita), extending from the deep reddish brown hypothecium (leptoclinoides-brown, textura intricata); outer excipular hyphae short-celled, cells angular, distinctly swollen
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Fig. 4: Buellia nashii.—A. Subsquamulose thallus with sessile apothecia (Nash 6724a).—
B. Close-up of thallus (Nash 6724a).—C. Cross section of an apothecium with dispersa-type exciple (Nash /5313b).—D. Overmature, slightly constricted ascospore with reduced septum thickening (Nash /53/3b).—E. Microrugulate ascospore ornamentation visible at a late stage of the spore ontogeny (Nash /5313b).—F. Conidia (Nash /53/3b).
(textura angularis) and carbonized with various amounts of a brown pigment (cf. elachista-brown); small amounts of a diffuse aeruginose pigment present but restricted to the outermost exciple cells (cinereorufa-green, HNO,+ violet); only the brown pigmentation continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown).
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ASCI—%-spored, clavate, Bacidia-type. ASCOSPORES (Fig. 4 D, E)—oblong to ellipsoid, rarely constricted with age, with obtuse ends, not curved, (11.0—)12.9-[14.5]- 16.1(-19.0) x (6.0—)6.5-[7.3]-8.1(—9.0) wm (n = 60); one-septate, proper septum briefly thickened but very soon becoming reduced during spore ontogeny, lateral wall thickenings absent (+ Physconia-type); ornamentation microrugulate.
PYCNIDIA—Rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980 and Vobis & Hawksworth 1981); conidia (Fig. 4F) simple, bacilliform, 5.0-10.0x 1.5—2.0 wm (n = 20).
CHEMISTRY—Typically with the depside atranorin and the depsidones norstictic (forming orange, needle-shaped crystals in the compound microscope) and connorstictic acid. Some specimens do not contain norstictic acid but instead are characterized by stictic and hypositictic acid. Specimens typically react K+ yellow to red (sometimes weak), P+ yellow (sometimes weak), C—, KC—, CK—. UV- (pale). The thallus is not amyloid, apothecia react amyloid in Lugol’s (always test with concentrated Lugol’s iodine or in the compound microscope; positive reactions can be very weak!).
SUBSTRATE AND ECOLOGY—0On a variety of siliceous (HCI-) rock substrates, rarely also on sandstones with small amounts of carbonates (HCI+).
DISTRIBUTION (Fig. 6)—The species is widely distributed throughout the Sonoran Desert Region and adjacent areas, extending into NE Mexico, but absent from low desert elevations.
NOTES—Thalli of B. nashii are very similar and almost as variable as those of B. dispersa (see the description of morphotype I in Bungartz et al. 2002). The two species nevertheless are reliably distinguished by their chemistry (2’-O-methylperlatolic acid in B. dispersa, vs. norstictic and/or stictic acid in B. nashii) and their different exciple pigmentation. No specimen with the chemistry of B. dispersa contains any traces of the aeruginose, HNO,+ violet pigment cinereorufa-green. In specimens of B. nashii this pigment is always present but usually confined only to the outermost cells of the exciple. Specimens should always be tested with HNO, for a reliable identification. Most specimens of B. nashii contain norstictic acid and are thus easily distinguished form B. dispersa because of the formation of distinct orange, needle-shaped crystals if KOH is applied to apothecial or thallus sections.
REPRESENTATIVE SPECIMENS EXAMINED—MEXICO. Baya Catirornia. Nash 26285, 26320b (ASU). Curauanua. Nash 13817, 13766 (ASU). Coanuma. Nash 6742c (ASU).—UNITED STATES. Arizona. Cochise Co. Weber $8837 (COLO). Coconino Co. Boykin 2849, 2674, Nash 30677, 4980 (ASU). Gila Co. Nash 7386 (ASU). Graham Co. Nash 766/ (ASU). Maricopa Co. Nash 84/1, 6266, 9850, 6659, 987, 5011 (ASU). Mohave
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Co. Nash 7303 (ASU). Pinal Co. Nash 6/5/ (ASU). Santa Cruz Co. Nash 3923] (ASU). Yavapai Co. Nash 34/37, 34149, 34188, 6361, Ryan 26958 (ASU). CaALtrornia. Inyo Co. Ryan /488/ (ASU). San Diego Co. Nash 7052 (ASU). CoLorapo. San Juan Co. Nash 17757 (ASU). Nevapa. Lincoln Co. Ryan 1/5862 (ASU). New Mexico. San Juan Co. Nash 16397, 16395, Rankert //7 (ASU). Uran. Kane Co. Nash 507], 5457, Nash 66/0 (ASU). San Juan Co. Nebecker 2800 (ASU). Washington Co. Nash /53/5 (ASU).
Buellia navajoensis Bungartz sp. nov. FIGURE 5
Thallus saxicolus, crustaceus, areolatus vel sublobatus, crassus, eburneus, sine hypothallo. Apothecia immersia vel adnatia, lecideina, marginibus propriis tenuis. Excipulum tenuis, fulvo-caeruleum, pigmentum aeruginosum continens, carbonaceum. Asci 8-spori. Sporae uniseptae, oblongae vel anguste oblongae, septibus incrassatus 13—25 x 6-10 ym. Pycnidia urceolata vel globosa. Conidia bacilliformes, 3-4.5 x 1-1.5 jum. Thallus acida norstictica et
connorstictica et substancia xanthonia continens. Medulla amyloideus.
Etymology: The species is named after the Native American Navajos and the type specimens
were collected on Navajo Sandstone.
Type: U.S.A. UTAH. Kane Co. About 16 km east of Kanab along U.S. Highway 89, in Johnson Canyon near the Kanab Stake Welfare Farm, ca. 1734 m, 37°02’14”N, 112°21°53”W, on Navajo Sandstone, Nebecker 1577 (ASU! — holotype designated here).
THALLUS (Fig. 5A, B)—Crustose, thick, + continuous, epilithic; areolate to sublobate often forming rosettes; prothallus absent; thallus surface matt and dull, not shiny, usually ivory, beige, rarely with a pinkish tinge, usually strongly pruinose, rarely weak or absent, phenocorticate; entire thallus filled with an abundance of calcium oxalate (H,SO,+ forming needle shaped crystals).
APOTHECIA—Lecideine, (0.3—)0.5—1.1(—1.4) mm in diameter, immersed, becoming adnate with age; proper margin black, thin, + persistent, excluded with age; disc black, usually with a fine white pruina, plane, becoming slightly convex with age; exciple narrow, poorly differentiated, of the aethalea-type (Fig. 5C) sensu Scheidegger (1993), i.e. inner excipular hyphae narrow, hyaline, prosoplectenchymatous (textura oblita), often reduced, similar in structure and orientation to the paraphyses, transient with the deep reddish brown hypothecium (leptoclinoides-brown, textura intricata), outer excipular hyphae parallel, moderately swollen (textura oblita) and moderately carbonized with various amounts of brown and aeruginose pigments (cf. elachista- brown & cinereorufa-green), pigmentation + continuous with the epihymenium; hymenium hyaline, not inspersed; paraphyses simple to moderately branched, apically swollen, with a brown pigment cap (cf. elachista-brown) and a diffuse, aeruginose pigment (HNO,+ violet, cinereorufa-green).
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ASCI—8-spored, clavate, Bacidia-type. ASCOSPORES (Fig. 5E-—I)—oblong to narrowly oblong, very rarely constricted, with obtuse ends, not curved, (13.0—)16.3— [18.6]—20.9(-25.0) x (6.0—)6.2-[7.1]-8.0(-10.0) wm (n = 60); one-septate, proper septum briefly thickened but very soon becoming reduced during spore ontogeny, lateral wall thickenings absent (+ Physconia-type); ornamentation microrugulate (to faintly striate); septum with septal pore canal, simple pore and undifferentiated pore plug; spore wall (Fig. 5G) differentiated into smooth to indistinctly cracked, thick perispore (0.30—0.40 ym), narrow intermediate layer (< 0.03 ym), thick proper spore wall (0.35—0.53 wm) and moderately thickened endospore (0.18—0.30 pm).
PYCNIDIA—Rare, urceolate to globose, unilocular, at maturity almost entirely occupied by densely branched conidiophores; conidiogeneous cells mostly terminal, rarely also intercalary (cf. conidiophore-type V sensu Vobis 1980); pycnidial ontogeny similar to the Umbilicaria-type (sensu Vobis 1980 and Vobis & Hawksworth 1981); conidia simple, bacilliform, 3.0—-4.5x 1.0—1.5 wm (n = 20).
CHEMISTRY—Typically with the depside atranorin and traces of the depsidone norstictic acid. Several xanthones have also been detected with HPLC by J. Elix: 4, 5- dichloronorlichexanthone, arthothelin, thiophanic acid. However, the thallus does not react C+ orange and the UV-fluorescence is only faint, possibly obscured by the large amount of Ca-oxalate pruina on the thallus surface. Thalli usually react K+ yellow, rarely K+ yellow-red (forming orange, needle-shaped crystals if observed in the compound microscope), P+ yellow, C— , KC—, CK—. UV+ pale yellow to beige. Thallus and apothecia react strongly amyloid.
SUBSTRATE AND ECOLOGY—0On limestone (HCI+ strongly reacting) or sandstone with some traces of carbonates (HCI+ weakly reacting, rarely HCI-).
DISTRIBUTION (Fig. 6)—Currently known only from the U.S.A. (Utah, New Mexico, Colorado, Northern Arizona, California).
NOTES—The thick, pruinose, areolate to sublobate, deeply amyloid thalli of B.
—>
Fig. 5: Buellia navajoensis.— A. Overview of areolate thallus (Nebecker /58/).—B. Close- up of areolate thallus (Nebecker /58/).—C. Cross section of an apothecium with aethalea- type exciple (Nebecker /58/).—D. Hymenium with asci, paraphyses and one ascospore (Nash /58/).—E. Mature ascospore: an indistinct septum can be seen in some of the spores (Nash /58/).—F. Immature ascospore prior to septum formation (Nash 1577 — holotype).— G. Mature ascospore (Nash /577 — holotype).—H. Wall layers in an immature ascospore: (1) outer layer (perispore and mucilaginous sheath not distinctly differentiated); (2) proper spore wall and (3) endospore beginning to become differentiated (Nash 1577 — holotype).—I. Wall layers in a mature ascospore: (as) ascus wall; (s) mucilaginous sheath; (1) perispore; (2)
intermediate layer; (3) proper spore wall; (4) endospore (Nash 1577 — holotype).
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