Overview of the rust fungi (Uredinales) occurring on Rosaceae in Europe

Stephan Helfer

Rose rust is caused by members of the genus Phragmidium, four of which are: P. mucronatum (Pers.) Schlect., P. tuberculatum Mull., P. fusiforme Schroet. and P. rosae-pimpinellifoliae (Rabh.) Diet. In the present study, preliminary identification of Phragmidium species was based on host range. Identification was then supported by measuring the dimensions of urediospores and teliospores. Axenic cultures of P. tuberculatum were initiated on culture media but these cultures were unable to reinfect rose leaves, therefore cultures of P. tuberculatum on leaf discs of 'Mme. Grégoire Staechelin' were initiated from urediospores. The infectivity of dry spores and spore suspensions of urediospores of P. tuberculatum directly from host tissue did not differ significantly; however, spore density of 30 × 10 4 per ml proved to be more infective than higher or lower densities. Surface-sterilized detached leaves showed significantly lower infection rates than leaves that were not surface sterilized. The urediospores on the leaf discs, survived when stored at 4°C for four weeks, but when stored for 12 weeks at the same temperature, they lost their viability and were not able to infect leaf discs.

This paper presents a checklist of rust fungi on family Poaceae in Pakistan together with their known host plants. A total of 80 taxa are mentioned here including 65 species of genus Puccinia and 15 of Uromyces. A host index is also provided.

Nova Hedwigia 81 3—4 325—370 Stuttgart, November 2005 Overview of the rust fungi (Uredinales) occurring on Rosaceae in Europe by Stephan Helfer Royal Botanic Garden Edinburgh, Edinburgh, EH3 5LR Scotland UK, Fax 0044 131 248 2901 With 3 figures Helfer, S. (2005): Overview of the rust fungi (Uredinales) occurring on Rosaceae in Europe. - Nova Hedwigia 81: 325-370. Abstract: The rust fungi (Uredinales) occurring on members of the rose family (Rosaceae) in Europe are described with particular emphasis on their host range and distribution. Rusts have been reported on less than half (46%) of the species of European Rosaceae with some genera being completely rust free. Other genera display a mixture of susceptible and immune species (e.g. Rubus: 56 susceptible / 20 immune; Alchemilla: 30 susceptible / 88 immune). Some rust genera are restricted to a single host species, whereas others parasitise many, mainly closely related hosts. Keys to the rust genera and species are provided, and all 41 rust species are described in detail. An index of host species is appended. Key words: Rose family diseases, taxonomy, host range, distribution. Zusammenfassung: Die Rostpilze (Uredinales) auf Rosaceen in Europa werden unter besonderer Berücksichtigung ihrer Verbreitung und ihrer Wirtspflanzen beschrieben. Roste sind von weniger als der Hälfte der Arten europäischer Rosaceen beschrieben, und einige Gattungen sind vollkommen befallsfrei. Andere Gattungen weisen teils anfällige teils immune Arten auf (z. B. Rubus: 56 anfällig / 20 immun; Alchemilla: 30 anfällig / 88 immun). Manche Rostgattungen sind auf eine einzige Wirtsart beschränkt, während andere viele, überwiegend nahe verwandte Wirtsarten befallen. Bestimmungsschlüssel zu den Rostgattungen und -arten sind angeführt, und alle 41 Rosaceenroste werden im Detail beschrieben. Ein Index der Wirtspflanzenarten ist angefügt. Stichwörter: Rosaceenkrankheiten, Taxonomie, Wirtswahl, Verbreitung. Introduction The family Rosaceae is represented in Europe by at least 473 species in 30 native genera (Valentine & Chatter 1968). It includes both herbaceous and woody plants as well as economically important fruit and ornamental crop species. It is of worldwide DOI: 10.1127/0029-5035/2005/0081-0325 0029-5035/05/0081-0325 $ 11.50 © 2005 J. Cramer in der Gebrüder Borntraeger Verlagsbuchhandlung, D-14129 Berlin · D-70176 Stuttgart 325 distribution, centred in North Temperate regions (Rowley 1993). Representatives of this family can be found in all regions of Europe (see Appendix 1), from the Mediterranean to the arctic and from oceanic to tundra and alpine habitats. The rust species parasitising members of the rose family belong to a broad variety of taxonomic groups. They are placed in Chaconiaceae, Phragmidiaceae, Pucciniaceae, Pucciniastraceae, Raveneliaceae and Uropyxidaceae (after Cummins & Hiratsuka 2003). Whilst the species concept of some of the 20 compatible host genera is somewhat debatable (especially Rubus, Alchemilla and Potentilla), the rusts are relatively well defined and the species are generally accepted. However, there still seems to be some debate concerning the higher taxonomic ranking of some of the species, especially in Leucotelium, Ochropsora and Tranzschelia (Thirumalachar & Cummins 1948, Gäumann 1959, Cummins & Hiratsuka 1983, 2003). For this study, the host nomenclature and taxonomy were based on the treatment in Flora Europaea (Valentine & Chatter 1968), which is generally accepted by mycologists (e.g. Vanky 1994, Braun 1995). Where necessary, synonymies were resolved using Kerguélen (1999). This paper aims to present a base line account of known compatible relationships, as well as providing a key for identification of rust fungi in European Rosaceae. Whilst many specimens of the represented taxa have been examined, it is not intended as a taxonomic revision of the species involved. Rather it represents current floristic data of host - pathogen interactions. It is hoped that this paper will stimulate the collecting and research of rosaceous rusts and further distribution data will become available in future. Materials and methods Herbarium (BERN, BPI, BRA, E, K, PRC, PUR, Z, ZT) specimens of most of the species and fresh specimens of selected species were examined using light microscopy and scanning electron microscopy. For light microscopy sori were dissected off the infected tissue and mounted in lactophenol-cotton blue for examination. SEM specimens were mounted on aluminium stubs using adhesive carbon tabs, sputter coated with gold/palladium for two minutes resulting in a deposition of ca 12nm, and examined in a Zeiss DSM 962 scanning electron microscope. Alternatively, specimens were examined uncoated at high vacuum or variable pressure (30 - 70 Pa) in a LEO 55 SEM. Specimens were determined using existing keys (Gäumann 1959, Wilson & Henderson 1966, Kern 1973, Gjærum 1974, Majewski 1977, 1979, and Minkevicius & Ignataviciute 1991) and original diagnoses. New species descriptions and keys were constructed using all of these data. European distribution data were extracted from herbarium data, distribution lists and local floras (above and Brandenburger 1994, Braun 1982, Denchev 1995, Farr et al. 2004, Guyot 1957, Henderson 2000, Karatygin et al. 2003, Kuprevich & Tranzschel 1957, Poelt 1985, Ryzhkind & Levkina 2004, Spooner & Butterfill 1999, Zwetko 2000). Host distribution data were verified using the Flora Europaea database (Anon 2004a). The morphological spore concept was applied (Laundon 1967, 1972) for rust life cycles. Spore characters were used as in Helfer (1992), with spine densities measured as units per area. Geographical codes were employed as in Flora Europaea (updated to the current political situation). They are as follows (Flora Europaea equivalent in brackets[] where updated): 326 Al Au Az Ba Be Bk Bl Br Bu Co Cr CS Da Fa Fe Ga Ge Gr Hb He Ho Hs Hu Is It Kr Lu Albania Austria with Liechtenstein Açores Baltic states: Estonia, Latvia, Lithuania, Kaliningradskaja Oblast’ of Russia [Rs(B)] Belgium Balkan states of Bosnia, Croatia, Herzegovina, Macedonia and Slovenia [Ju] Islas Baleares Britain, including Orkney, Zetland and Isle of Man; excluding Channel Islands and Northern Ireland Bulgaria Corse Kriti (Creta) with Karpathos, Kasos and Gavdhos Czech Republic and Slovakia [Cz] Denmark Færöer Finland (Fennia), including Ahvenanmaa (Åland Islands) France (Gallia), with the Channel Islands (Îles Normandes) and Monaco; excluding Corse Germany Greece, excluding those islands included under Kriti (supra) and those which are outside Europe as defined for Flora Europaea Ireland (Hibernia); both the Republic of Ireland and Northern Ireland Switzerland (Helvetia) Netherlands (Hollandia) Spain (Hispania) with Gibraltar and Andora; excluding Islas Baleares Hungary Iceland (Islandia) Italy, including the Arcipelago Toscano; excluding Sardegna and Sicilia Crimea (Krym) [Rs(K)] Portugal (Lusitania) No Norway Po Poland Rm Romania RB Belarus and Central Russia [Rs(C)] RE South-eastern Russia: Lower Don, Lower Volga, Transvolga [Rs(E)] RN Northern Russia: Karelo-Lapland, Dvina-Pecora [Rs(N)] Sa Sardegna Sb Svalbard, comprising Spitsbergen, Björnöya (Bear Island) and Jan Mayen Si Sicilia, with Pantelleria, Isole Pelagie, Isole Lipari and Ustica; also the Malta archipelago Su Tu UM Sweden (Suecia), including Öland and Gotland Turkey (European part), including Gökçeada (Imroz) Moldova and Ukraine, excluding Crimea, [Rs(W)] 327 Figure 1: Recorded susceptibility to rust fungi of the genera of native European Rosaceae. All species recognised in Flora Europaea were included. Results and discussion There are 41 rust species reported on a total of 216 species (out of 474, sensu Valentine & Chatter 1968, excluding naturalised aliens and related species) of Rosaceae in Europe resulting in at least 408 disease relationships. The rust species belong to the following 13 genera (number of species in brackets): Frommeella (1) Gymnoconia (1) Gymnosporangium (9) Kuehneola (1) Leucotelium (1) Ochropsora (1) Phragmidium (15) Puccinia (2) Pucciniastrum (3) Trachyspora (2) Tranzschelia (2) Triphragmium (2) Xenodochus (1) The distribution on susceptible host species among the genera of the Rosaceae is presented in Fig. 1. Several small genera exhibit a complete lack of rust parasites. These comprise ten genera, five of which are monotypic in Europe. Spiraea is the largest genus of this group with 13 species in Europe (however, five of these 328 Figure 2: Host specificity of the rust fungi infecting native European Rosaceae. are not native European). Farr et al. (2004) report no rust species among the many fungal pathogens of this genus worldwide. Other genera contain a large proportion of host species (e.g. Rubus with 56 out of 76 species [including R. fruticosus] as reported host plants = 74%); conversely some larger genera (e.g. Alchemilla) have only few host plants (30 of 118 = 25%). A total of 257 species (54%) have so far not been reported to host rust species. Whilst this may to some extent reflect problems of nomenclature in these genera or the low frequency of the species involved, there appears to be a biased distribution which may be interpreted as exhibiting patterns of natural susceptibility or resistance. As far as the rust parasites are concerned there is great variation in host specificity (Fig. 2) with the genus Phragmidium (represented by 15 species) parasitising 122 species of the Rosaceae and P. violaceum alone having a host range of 45 species (all of them Rubus spp.). Holm (1980) and Durrieu (1987) discussed the possible affinities of the rosaceous rusts and their host taxa and endeavoured an explanation of their evolution and their distribution. Through lack of herbarium specimen data or published records, geographical distribution data of rosaceous rusts were unsatisfactory for the following areas: Al, Be, Bl, Co, Cr, Ho, Sa, Sb, Si and Tu. 329 Key to the genera A.) using teliospore characters: 1 Telia forming subepidermal crusts or formed within epidermal cells, teliospores sessile......... 2 Telia erumpent or +/- pulverulent, teliospores on pedice......................................................... l3 2 (1) Telia dark brown-black. Teliospores persistently subepidermal or intra-epidermal, divided into 2-4 cells by anticlinal septa.................................................................. Pucciniastrum Telia pale or colourless. Teliospores initially covered by epidermis, single celled, probasidial................................................................................................................. Ochropsora 3 (1) Teliospores 1-celled............................................................................................. Trachyspora Teliospores consisting of more than one cell........................................................................... 4 4 (3) Teliospores 2-celled............................................................................................................... 5 Teliospores consisting of more than two cells........................................................................ 10 5 (4) Telia often formed in early spring on gymnosperms, erumpent as cushions, crests or “horns”, teliospores smooth.................................................................................... Gymnosporangium Telia never erumpent.............................................................................................................. 6 6 (5) Teliospore surface verrucose (Fig. 3C) deeply constricted andcells easily separating................... ...................................................................................................................... Tranzschelia Teliospore surface smooth or rugose to pseudoreticulate (Fig. 3H), cells not easily separating ........................................................................................................................................ 7 7 (6) Teliospore surface rugose to pseudoreticulate (Fig. 3H)............................... Puccinia sieveriae Teliospore surface smooth...................................................................................................... 8 8 (7) Teliospore with distinct papillae (Fig. 3A)............................................................. Gymnoconia Papillae lacking................................................................................................................ 9 9 (8) Basidia an apical elongation of telial cells................................................................ Leucotelium Promycelium germinating from +/- lateral, distinct pores in teliospore........................................ ............................................................................................................ Puccinia waldsteiniana 10 (4) Teliospores three-celled, triangular (Fig. 3D)................................................... Triphragmium Teliospores many-celled (1- 22, variable within sorus)......................................................... 11 11 (10) Teliospores cylindrical, articulated, wall colourless, germpores projecting............... Kuehneola Teliospores +/- curved, wall brown, germpores not projecting............................................. 12 12 (11) All teliospore cells with 1 germpore...................................................................... Frommeella Most teliospore cells with two or more germpores................................................................. 13 13 (12) All teliospore cells (including apical cell) with two or more germpores............... Phragmidium Teliospore cells with 2 or more germpores except apical cell (1 germpore)............ Xenodochus B.) key, using other than teliospore characters: 1 Rust (any spore stage) on Filipendula................................................................ Triphragmium Rust (any spore stage) on Alchemilla.................................................................... Trachyspora Rust (II and III) on Geum or Waldsteinia..................................................................... Puccinia Rust (II and III) on Agrimonia......................................................... Pucciniastrum agrimoniae Rust (0 and I) on subfamily Pomoideae...................................................... Gymnosporangium Rust on other rosaceous genera or no spermogonia and aecia on Pomoideae........................... 2 2 (1) Uredinia never present............................................................................................................ 3 Uredinia normally present....................................................................................................... 4 3 (2) Aecia small and semi systemic, on Rubus............................................... Gymnoconia peckiana Aecia large (to 10 mm), on Sanguisorba........................................... Xenodochus carbonarius 330 4 (2) Uredinial paraphyses present................................................................................................. 5 Uredinial paraphyses absent................................................................................................... 7 5 (4) On Rosa or Rubus.............................................................................................. Phragmidium On Prunus, urediniospores smooth at apex, paraphyses dispersed..................... Tranzschelia On Pomoideae, Aruncus or Prunus, urediniospores evenly echinulate, uredinial paraphyses peripheral, forming small circular cups........................................................ Ochropsora ariae On Potentilla.......................................................................................................................... 6 6 (4) Aecia uredinioid, urediniospores very densely echinulate (0.8 - 1.1 spines/µm2)........................ .......................................................................................................... Frommeella tormentillae Aecia caemoid, urediniospores more distantly echinulate (0.46 - 0.58 spines/µm2) or verrucose .................................................................................................................... Phragmidium 7 (4) On Sanguisorba, aecia ca. 1mm.................................................... Phragmidium sanguisorbae On Prunus............................................................................................................................. 8 On Rubus............................................................................................................................... 9 8 (7) Urediniospores 17-30 × 15-20 µm, wall brown.......................................... Leucotelium cerasi Urediniospores 15-24 × 10-16 µm, wall hyaline............................... Pucciniastrum areolatum 9 (7) Uredinia pulverulent, occasionally covering calyx and stem...................... Kuehneola uredinis Uredinia covered by epidermis, peridial, semi-systemic................. Pucciniastrum arcticum Account of rust species: Frommeella Cummins & Y. Hirats.: This small genus is closely related to Phragmidium and differentiated by the single germ pore per teliospore cell of Frommeella compared with 2-4 germ pores in Phragmidium. Originally the species were allocated to the genus Frommea Arthur. However, according to Laundon (1975) the genus Frommea can be synonymised with Phragmidium, and therefore a new generic name had to be found. Viennot-Bourgin (1954) reported another rust of this genus on Duchesnea indica Focke in the Pyrenees: Frommeella mexicana var. indicae J.W. McCain & J.F. Hennen, however, as the host is not a native European, I have not included it in the formal account. 1. Frommeella tormentillae (Fuckel) Cummins & Y. Hirats. (1983). SYNONYMS: Phragmidium tormentillae Fuckel (1869); Frommea obtusa (F.Strauss) Arthur (1917). Gäumann (1959): 1174; Wilson & Henderson (1966): 110; Gjærum (1974): 40; Majewski (1977): 298; Minkevicius & Ignataviciute (1991): 189; Frommea tormentillae (Fuckel) U.Braun (1978): Braun (1982): 228; Poelt (1985): 58 (as Frommea potentillae (Fuckel) U.Braun). Spermogonia: epiphyllous, intraepidermal (type 10), in small groups in reddish slightly hypertrophied spots, inconspicuous. Uredinia I: epiphyllous, subepidermal in origin; surrounding spermogonia. Spores: borne singly; 18-28 × 12-18 µm; wall thickness: 1-1.5 µm; orange yellow when fresh; walls echinulate above nearly smooth below, germpores obscure. Uredinia II: hypophyllous; small (0.5 mm); light yellow; initially covered by epidermis; with occasional paraphyses 7-9 µm thick. Spores: 16-25 × 13-20 µm; wall thickness: 1-1.5 µm; pale yellow; finely echinulate (0.8-1.1 spines/µm2); germpores 3-4, obscure. 331 332 Telia: hypophyllous; small (0.5 mm); light brown (cinnamon). Spores: 2-7 celled; 40-150 µm (depending on cell number) × 18-24 µm, cylindroid to clavoid, often curved or ‘bent’; slightly constricted where cells meet; wall smooth colourless to pale brown; thin at base (1 µm) thickening towards apex (8 µm); germpores 1 per cell at apical cell end; cell contents orange; pedicels of varying length, 7-10 µm thick, persistent. Life cycle: autoecious; brachycyclic. Hosts: Potentilla erecta (L.) Räuschel; P. anglica Laicharding, P. recta L., P. reptans L. Distribution: Europe: Au, Az, Br, CS, Da, Fe, Ge, Hb, He, Hs, Hu, It, No, RE, Rm, RN, UM; Asia, North America. Uncommon. Gymnoconia Lagerh.: A monotypic genus with unique teliospore morphology. According to Laundon (1975), a new genus had to be established to cover the autopsis form found on rosaceous hosts. Consequently, Cummins & Hiratsuka (1983) published the new combination under a new generic name: Arthuriomyces. The old name Gymnoconia was thought to apply to the microcyclic form only. However, Cummins & Hiratsuka (2003) point out that, when the genus was first described, it would have occurred as autopsis form, making the change superfluous. Only four host species have been confirmed for this rust from subgenus Cyclactis and Rubus, section Rubus subsections Discolores and Appendiculati. There are many further species in these taxa which may have been overlooked as hosts. 2. Gymnoconia peckiana (Howe) Trotter SYNONYMS: G. interstitiale (Schltdl.) Lagerh. (1894); Puccinia peckiana Howe (1873); Arthuriomyces peckianus (Howe) Cummins & Y. Hirats. (1983). Gäumann (1959): 209; Gjærum (1974): 42; Majewski (1977): 294; Poelt (1985): 58; Minkevicius & Ignataviciute (1991): 111. IMI 2011 . Fig. 3A. Spermogonia: on both sides of leaf, subcuticular, protruding cone shaped to 75 µm. Aecia: hypophyllous, caeomoid, 0.5-1.5 mm, covering the whole surface of the leaf; initially covered by epidermis, later surrounded by remainders of the epidermis; orange yellow. Spores: ellipsoid 19-38 × 16-34 µm; wall thickness: 1.5-2 µm; colourless; germpores indistinct; verrucose, 1.5-2 µm; spore contents orange. 1 International Mycological Institute: Descriptions of fungi and bacteria. (earlier: Description of pathogenic fungi and bacteria); number of description. Figure 3: Scanning electron micrographs of European rosaceous rust fungi; A. Gymnoconia peckiana (E92371) on Rubus arcticus, teliospores; length of bar: 10 µm. B. Phragmidium bulbosum (E192370) on Rubus caesius, urediniospores; length of bar: 10 µm. C. Tranzschelia discolour (E SH 1009) on Prunus domestica, teliospores; length of bar: 20 µm. D. Triphragmium ulmariae (E H410) on Filipendula ulmaria, teliospores; length of bar: 10 µm. E. Phragmidium acuminatum (E192372) on Rubus saxatilis, teliospores; length of bar: 10 µm. F. Gymnosporangium fusisporum (Z 21) on Cotoneaster integerrimus, aeciospore ornamentation; length of bar: 1µm. G. Trachyspora melospora (ZT 16) on Alchemilla hoppeana, teliospores; length of bar: 10 µm. H. Puccinia sieversiae (PRC 4) on Geum reptans, teliospores; length of bar: 10 µm. 333 Uredinia: lacking. Telia: hypophyllous; small (0.3 mm); on yellow spots; dark brown; partly covered by epidermis. Spores: 2-celled; 30-50 × 18-30 µm; polymorphic; lower cell often tapering where porus appears; wall thin (1-1.5 µm); often with 1-6 papillae around porus; germpores 1 in each cell; pedicels colourless; spores readily detaching. Fig. 3A. Life cycle: autoecious; demicyclic. Hosts: Rubus arcticus L., R. chloocladus W.C.R. Watson, R. serpens Weihe ex Lej. (as R. hispidissimus Sudre), R. saxatilis L.. Distribution: reported in Europe from: Br, Fe, Ge, Hu, Is, No, RN, Su. Gymnosporangium R. Hedwig: With nine species in Europe, this genus represents heteroecious, demicyclic (lacking uredinia) rusts with rosaceaous plants of the subfamily Maloideae as alternate hosts and species of Juniperus (in Europe) as telial hosts. Key to the species: 1 Spore stage III present (on Juniperus or Cupressus2 )........................................................... 2 Spore stages 0 and I present (on Rosaceae)........................................................................... 10 2 (1) Teliospores elongate, spindle shaped, 4-5 times as long as broad............................................. 3 Teliospores oval, not more than twice as long as broad............................................................. 5 3 (2) Teliospores with single germ pore per cell, telia caulicolous............................... G. fusisporum Teliospores with two germpores near septum.......................................................................... 4 4 (3) Teliospores broad (20-34 µm)......................................................................... G. clavariiforme Teliospores less than 18 µm broad, telia caulicolous, forming “witches’ brooms”... G. gracile 5 (2) Teliospores papillate (3-6 µm), telia foliicolous................................. G. torminali-juniperinum Not so................................................................................................................................... 6 6 (5) Teliospore germpores apical.......................................................................... G. amelanchieris Teliospore germpores near septum.......................................................................................... 7 7 (6) Telia conical, laterally compressed........................................................................ G. confusum Telia not conical, either tremelloid or on fusiform swellings.................................................... 8 8 (7) Telia applanate, becoming tremelloid when wet................................................ G. tremelloides Telia on fusiform swellings..................................................................................................... 9 9 (8) Telia 8-10 mm (10-20 mm when wet)...................................................................... G. sabinae Telia 1-3 mm, flat.................................................................................................. G. cornutum 10 (1) Peridia short (up to 2 mm)..................................................................................................... 11 Peridia longer 3-5 mm........................................................................................................... 13 11(10) Aeciospores large (30-50 × 25-35 µm) aecia foliicolous.................................... G. tremelloides Aeciospores smaller (19-29 × 16-25 µm), aecia foliicolous, caulicolous or fructicolous........ 12 12 (11) Peridial cells clearly ridged, 60-90 µm long........................................................... G. confusum Peridial cells verrucose with roundish papillae, 80-120 µm long.............................. G. gracile 2 Gymnosporangium minus Crowell is the only species found on Cupressus in Europe. No rosaceous hosts are known for this species. The American G. cupressi Long & Godding is the most similar heteroecious species on Cupressus and alternates with Amelanchier spp. (Kern 1973). 334 13 (11) Peridia balanoid (not dehiscent at apex)................................................................... G. sabinae Peridia soon dehiscent.......................................................................................................... 14 14 (13) Aecia on leaf galls (clearly swollen leaf tissue).............................................. G. amelanchieris Aecia not on galls................................................................................................................. 15 15 (14) Aecia only foliicolous...................................................................... G. torminali-juniperinum Aecia foliicolous, caulicolous or fructicolous....................................................................... 16 16 (15) Peridial cells with ridges of varying length......................................................... G. fusisporum Peridial cells with papillae of varying size............................................................................. 17 17 (16) Peridial papillae roundish of varying size........................................................ G. clavariiforme Peridial papillae short, delicate.............................................................................. G. cornutum 3. Gymnosporangium amelanchieris E.Fisch. (1911). SYNONYMS: Gymnosporangium juniperinum (L.) Fr. f.sp. amelanchieris (E. Fisch.) Gäumann (1959). [Majewski (1977): 278;] Kern (1973): 25; Braun (1982): 228; Poelt (1985): 59. Gäumann (1959): 1172. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; first yellowish becoming black. Aecia: hypophyllous; roestelioid; on gall like protuberances; forming long peridia (3-5 mm); cornute, dehiscent at apex, becoming split along the sides; peridial cells rhomboid 29-39 µm thick, 80-100 µm long; outer wall 2-3 µm inner walls 7-9 µm thick. Spores: globoid; 18-23 × 25-29 µm; wall 1.5-2.5 µm thick; light brown; 6-10 pores; verrucose (approx. 1.1 warts/µm2); spore contents hyaline. Telia: caulicolous, on swellings, small on needles (1-2 mm), a little bigger on the stem (3-5 mm); applanate; brown. Spores: 2-celled; 19-32 × 35-55 µm; light brown; germpores 1-2 (apical). Hosts: 0&I: Amelanchier ovalis Medicus, Pyracantha coccinea M.J.Roemer; III: Juniperus L. sect. Oxycedrus. Distribution: In Europe: Au, Ga, Ge, He, Hs, Hu, It, Lu; Asia, Africa (North). This rust has been considered as a forma specialis of G. torminali-juniperinum (Kern 1973), from which it is different by having less strongly thickened peridial cell walls and moderately rugose ornamentation. Judging by the collections available this rust is often overlooked or has been included in G. torminali-juniperinum. 4. Gymnosporangium clavariiforme (Pers.) DC. SYNONYM: Tremella clavariaeformis Pers. (1805). Gäumann (1959): 1153 [as Gymnosporangium clavariaeforme (Jacq.) DC.]; Wilson & Henderson (1966): 116; Kern (1973): 31; Gjærum (1974): 44; Majewski (1977): 270; Braun (1982): 229; Poelt (1985): 59; Minkevicius & Ignataviciute (1991): 173. IMI 542. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous; roestelioid; 0.5 mm wide 2-3 mm long; tubular, soon lacerate; peridial cells rhomboid, long and narrow 80-130 µm. Spores: 18-23 × 25-29 µm; globoid; wall 1.5-3 µm thick; light brown; densely verrucose (1.1 warts/µm2); germpores 6-10. 335 Telia: on stems and needles; 5 mm by 2 mm thick, swelling when wet; elongate on swellings of the branches; yellowish-dark brown. Spores: 2-celled; 20-34 × 50-110 µm; lanceolate or fusiform, wall thickness 1 µm in centre of sorus to 2.5 µm at sorus margin; light brown at margin pale in centre; slightly constricted at septum; germpores 2 near septum; pedicels cylindrical, very long; persistent. Hosts: 0&I: Amelanchier ovalis Medicus, Crataegus azarolus L., Cr. calycina Peterm., Cr. laciniata Ucria, Cr. laevigata (Poiret) DC., Cr. monogyna Jacq., Cr. nigra Waldst. & Kit., Cr. pentagyna Waldst. & Kit. ex Willd., Cr. sanguinea Pallas, Cydonia oblonga Miller, Malus domestica Borkh., Pyrus communis L., Sorbus aria (L.) Crantz, S. aucuparia L., S. latifolia (Lam.) Pers., S. meinichii (Lindeb.) Hedl., S. minima (A. Ley) Hedl. (as S. arranensis Hedl.), S. neglecta Hedl., S. torminalis (L.) Crantz; III: Juniperus communis L. Distribution: reported in Europe from: Au, Ba, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Ho, Hu, It, Lu, No, Po, Rm, RN, Si, Su, UM; elsewhere in the northern hemisphere. Probably the most common species of the genus Gymnosporangium on non-cultivated host plants in Europe, this rust is very conspicuous with the production of bright orange-red lesions on a relatively large number of host species. The author found it also on bonsai plants (unpublished). This rust is a ‘founder member’ of the genus Gymnosporangium (Kern 1973). 5. Gymnosporangium confusum Plowr. (1889): Gäumann (1959): 1162; Wilson & Henderson (1966): 117; Kern (1973): 34; Gjærum (1974): 45; Majewski (1977): 272; Braun (1982): 229; Poelt (1985): 60; Minkevicius & Ignataviciute (1991): 175. IMI 544. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous and caulicolous; at times on fruits; roestelioid; 1-2 mm high; peridium quickly losing tubular form; lacerate nearly to base; in red spots; peridial cells with strong +/- longitudinal ridges and bumps, 60-90 µm long. Spores: 19-22 × 19-27 µm; globoid; wall 2-3 µm thick; light brown; finely verrucose [>1.2 spines/ µm2)]; germpores 6-10. Telia: caulicolous; on slight fusiform enlargements; 5-8 mm high; conical or laterally compressed; dark brown. Spores: on long pedicels; 20-26 × 35-45 µm; ellipsoid, at times narrower and longer; wall thickness: 2-3 µm or less; dark brown; germpores 2 near septum. Hosts: 0&I: Cotoneaster integerrimus Medicus, Co. nebrodensis (Guss.) K.Koch, Co. niger (Thunb.) Fries, Co. nummularia Fischer & C.A.Meyer, Crataegus azarolus L., Cr. laevigata (Poiret) DC., Cr. laciniata Ucria, Cr. monogyna Jacq., Cr. sanguinea Pallas, Cydonia oblonga Miller, Mespilus germanica L., Pyracantha coccinea M.J. Roemer, Pyrus communis L., Sorbus aucuparia L., S. latifolia (Lam.) Pers., S. torminalis (L.) Crantz; III: Juniperus L. sect. Oxycedrus and Sabina. 336 Distribution: Europe: Au, Be, Br, Bu, Cr, Da, Ga, Ge, Gr, He, Hs, Hu, Kr, Lu, No, Rm, Tu UM; Asia, Africa. The telial stage of this rust, similar to the previous species resembles that of G. sabinae, a fact which lead to its specific name. The aecial stages of the three fungi are readily distinguished by their peridial cell sculpturing (see key). 6. Gymnosporangium cornutum (Pers.) Arthur (1911). SYNONYMS: Gymnosporangium juniperinum (L.) Fr. p.p.. Gäumann (1959): 1170; Wilson & Henderson (1966): 121; Kern (1973): 37; Gjærum (1974): 46; Majewski (1977): 276; Poelt (1985): 60. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous at times on fruits; roestelioid; 3-5 mm high; cornute; on yellow spots later turning reddish; peridial cells rhomboid; 60-110 µm long; inner and side walls 8-12 µm thick; moderately rugose. Spores: 16-25 × 21-29 µm; globoid; wall thickness: 2-2.5 µm; brown; finely verrucose (1.1 spines/µm2); germpores 6-10. Telia: mainly caulicolous; on fusiform swellings of smaller branches; dark brown; occasionally foliicolous. Spores: 2-celled; 15-24 × 30-55 µm; ellipsoid; often narrowed above and below; wall thickness: 1-2 µm; brown; germpores 1-2 near septum. Hosts: 0&I: Sorbus aucuparia L., S. chamaemespilus (L.) Crantz, S. domestica L., S. hybrida L., S. intermedia (Ehrh.) Pers., S. latifolia (Lam.) Pers., S. mougeotii Soyer-Willemet & Godron; III: Juniperus L. sect. Oxycedrus. Distribution: Europe: Au, Ba, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, Is, It, Lu, No, Po, RE, Rm, RN, Su, UM; Asia; Africa; North America. Kern (1973) reports the occurrence of this rust on Malus (apparently rare). The author could not find confirmation for any occurrence on this host in Europe. 7. Gymnosporangium fusisporum E.Fisch. (1918). SYNONYM: G. confusum f.sp. fusisporum Gäumann (1959). Gäumann (1959): [1164]; Kern (1973): 50. Fig. 3F. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous; foliicolous rarely caulicolous or fructicolous; roestelioid; up to 5mm; conical to cylindrical; peridium soon rupturing at apex and becoming laciniate to base; peridial cells rhomboid rugose to verrucose. Spores: 18-24 × 19-35 µm; globose or broadly ellipsoid; wall thickness: 2-3 µm; light brown; finely verrucose [1.2 warts/µm²], Fig. 3F; germpores 6-9. Telia: caulicolous on fusiform swellings; size varying; conical or laterally compressed; dark brown. Spores: 2 celled; 16-24 × 46-90 µm; fusiform sometimes falcate; wall thickness: 1.5-2.5 µm; germpores: 1 near septum. 337 Hosts: 0&I: Cotoneaster integerrimus Medicus, C. nebrodensis (Guss.) C. Koch; III: Juniperus L. sect. Sabina. Distribution: Europe: He, Kr, UM; Africa. This rust is very similar to G. confusum in its aecial stage (but see aecial size), however, their telial stage (especially the teliospores) digress and justify the differentiation. 8. Gymnosporangium gracile Pat. (1902). Gäumann (1959): 1156; Kern (1973): 55; Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous; fructicolous or caulicolous; roestelioid; 0.4-1 mm; cylindrical; peridial cells long and narrow: 17-32 × 80-120 µm. Spores: 25-28 µm; globular; wall thickness: 2-3 µm; brown. Telia: caulicolous sometimes on swollen branches; usually forming witches’ brooms; 1-2 mm wide 3-4 mm high; cylindrical; orange. Spores: 2-celled; 12-18 × 50-90 µm; lanceolate or fusiform; wall thickness: 1-1.5 µm; light; germpores: 2 near septum; pedicels: extremely long. Hosts: O&I: Amelanchier ovalis Medicus, Crataegus monogyna Jacq., Cydonia oblonga Miller; III: Juniperus L. Distribution: southern Europe: Bu, Ga, Gr, Hs, It; Northern Africa; southern North America. This species has long been included in G. clavariiforme, from which it is indistinguishable in its aecial stage. Kern (1964), after considering evidence by other urediniologists, finally accepted it as separate species on account of the telia producing witches’ brooms on their Juniperus hosts. 9. Gymnosporangium sabinae (Dicks.) G.Winter (1884). SYNONYM: G. fuscum DC. (1805). Gäumann (1959): 1158; Wilson & Henderson (1966): [119]; Kern (1973): [49]; Gjærum (1974): 47; Majewski (1977): 274; Braun (1982): 229; Poelt (1985): [60]; Minkevicius & Ignataviciute (1991): 174. IMI 545. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous on galls; also caulicolous producing cankers; occasionally fructicolous; roestelioid; 0.5-1 mm diam. 2-5 mm high; peridium apex remaining bluntly conical, walls later bulging (balanoid); light; peridial cells rhomboid; 65100 × 20-28 µm; verrucose or papillose; walls 14-20 µm. Spores: 22-29 × 25-31 µm; globoid to ellipsoid; wall thickness: 3-4.5 µm; dark brown; finely verrucose [1.2 warts/µm²]; germpores 6-10. Telia: caulicolous on fusiform swellings; 8-10 mm; swelling to 10 × 20 mm when wet; at first pulvinate; later conical or laterally compressed; dark brown. Spores: 2338 celled; 18-30 × 40-50 µm; ellipsoid to oblong; wall thickness: 1-3 µm; nearly colourless to dark brown; germpores: 2 near septum; pedicels: hyaline; very long. Hosts: 0&I: Pyrus amygdaliformis Vill., P. communis L., P. elaeagrifolia Pallas, P. nivalis Jacq., P. pyraster (L.) Burgsd., P. salicifolia Pallas, P. syriaca Boiss. III: Juniperus communis L., J. sabina L. Distribution: Europe: Au, Ba, Br, Bk, Bu, CS, Da, Ga, Ge, Gr, He, Hs, Hu, It, Kr, Lu, No, Po, Rm, UM; Asia; Africa. Known also as the ‘Pear Rust’ this is one of the first (Puccinia graminis being the first) rusts for whom the heteroecious life cycle has been established by artificial inoculation (Oersted 1865). Arguably, this rust represents the most economically important parasite among the genus Gymnosporangium. It is extremely widespread where the two hosts live together. 10. Gymnosporangium torminali-juniperinum E. Fisch. (1911). SYNONYM: G. juniperinum (L.) Fr. f.sp. torminali-juniperinum E. Fisch. Gäumann (1959): [1172]; Kern (1973): 76; Poelt (1985): 61. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous; roestelioid; 2-5 mm high; peridium dehiscent at apex; becoming more or less split along the sides; light; peridial cells rhomboid; 65-100 µm long. Spores: 18-24 × 24-27 µm; broadly ellipsoid; wall thickness: 1.5-2.5 µm; cinnamon brown; finely verrucose (1.1 warts/µm2); germpores 6-10. Telia: foliicolous chiefly epiphyllous; 0.5-1.5 mm high; hemispherical. Spores: 2celled; 21-30 × 35-49 µm; broadly ellipsoid; wall thickness: 1-1.5 µm; brown; papilla on apex 3-6 µm. Hosts: 0&I: Sorbus aucuparia L., S. chamaemespilus (L.) Crantz, S. domestica L., S. hybrida L., S. latifolia (Lam.) Pers., S. mougeotii Soyer-Willemet & Godron, S. torminalis (L.) Crantz; III: Juniperus L. Distribution: Europe: Au, Br, Ga, Ge, He, Hu, It, Lu, RE, Su; northern Africa. A fungus which is very similar to G. cornutum, but whose teliospores are sufficiently different to justify separation; also the peridial cells show differences (see key). 11. Gymnosporangium tremelloides R.L.Hartig (1882); Gäumann (1959): 1166; Kern (1973): 78; Gjærum (1974): 47; Majewski (1977): 279; Braun (1982): 229; Poelt (1985): 60; Minkevicius & Ignataviciute (1991): 171. IMI 549. Spermogonia: epiphyllous; deep seated; globoid in the centre of leaf discolourations; 110 µm; protruding as distinct cones (130-140 µm); first yellowish becoming dark brown. Aecia: hypophyllous; roestelioid; 0.5-1.5 mm high; cylindrical; on swollen spots; peridium becoming fimbriate to base; pale; peridial cells rhomboid; 31-35 × 60-90 µm; rugose with thickened inner and lateral walls. Spores: 25-35 × 30-50 µm; large; 339 very variable; globoid-ellipsoid; wall thickness: 3-5 µm; dark brown; finely verrucose [>1.2 warts/µm²]; germpores 6-8 dispersed. Telia: caulicolous on fusiform galls; 5 × 20 mm; 1-2 mm high; applanate; becoming tremelloid when wet; orange brown. Spores: 2-celled; 20-30 × 35-70 µm; ellipsoid; wall thickness: 1-2.5 µm; light brown to dark brown; smooth; germpores: 2 in lower cell near septum; 2-3 in upper cell (1-2 near the septum 1 at apex). Hosts: 0&I: Cydonia oblonga Miller, Malus domestica Borkh., M. sylvestris Miller, Sorbus aria (L.) Crantz, S. aucuparia L., S. chamaemespilus (L.) Crantz, S. latifolia (Lam.) Pers., S. torminalis (L.) Crantz; III: Juniperus L. sect. Oxycedrus. Distribution: Europe: Au, Ba, Bu, CS, Da, Fe, Ga, Ge, He, Hs, Hu, It, No, Rm, Su, UM; Asia; western Africa; North America. Uncommon. Unfortunately this rust has suffered under taxonomic confusion (Kern 1973), and its distribution may therefore not be as well established as could be wished. However, its exceptionally large aeciospores could help in correcting this calamity by providing a clear character for identification. Kuehneola Magnus: The genus Kuehneola is separated from Phragmidium by possessing single germ pores in the teliospore cells. In Europe it is represented by a single species on Rosaceae. 12. Kuehneola uredinis (Link) Arthur (1906). SYNONYMS: Oidium uredinis Link (1824); K. albida (J.G.Kühn) Magnus (1898). Gäumann (1959): [197]; Wilson & Henderson (1966): 108 [as Kuhneola uredinis (Link) Arthur]; Gjærum (1974): 52; Majewski (1977): 295; Braun (1982): 230; Poelt (1985): 61; Minkevicius & Ignataviciute (1991): 113. IMI 202. Spermogonia: epiphyllous; subcuticular; large, 150-200 µm; pustular; on reddish spots. Uredinia I: epiphyllous around spermogonia; small, 0.5 mm; round, arranged in circles at times confluent; orange yellow. Spores: borne singly on pedicels; 18-29 × 16-23; globoid to obovoid; wall thickness: 2 µm; colourless; finely verrucose; germpores obscure; contents: yellow. Uredinia II: hypophyllous at times on calyx and stem; small [0.1-0.5 mm]; round; lemon yellow; without paraphyses. Spores: 18-29 × 16-23; globoid to obovoid; wall thickness: 1.5-2 µm; colourless; finely verrucose-echinulate [0.8 warts/µm²]; germpores: 3-4 equatorial; indistinct. Telia: hypophyllous; 0.2-0.5 mm; singular or in groups, never coalescing; yellowish or whitish; soon uncovered. Spores: 2-12 in chains; individual spores 17-30 × 15-20 µm; trapezoid; wall thickness: basal 1.5-2 µm; apical up to 4 µm; pale; germpores: apically lateral in each spore; forming small projections; pedicels: thin walled; non-persistent. Life cycle: autoecious; brachycyclic. Hosts: Rubus affinis Weihe & Nees [also as R. fissus Lindl.], R. amoenus Koehler, R. arrhenii Lange, R. caesius L., Rubus candicans Weihe ex Reichenb. [as R. 340 arduennensis Lej.], R. canescens DC., R. chloocladus W.C.R.Watson [as R. vestii Focke], R. divaricatus P.J.Müller, R. fruticosus L. s.l., R. fuscater Weihe & Nees [as R. adornatus P.J.Mueller ex Wirtgen], R. fuscus Weihe & Nees, R. glandulosus Weihe ex Lej. & Court., R. gratus Focke [also as R. sciocharis (Sudre) W.C.R.Watson], R. gremlii Focke [also as R. ferox Vest ex Tratt.], R. hirtus Waldst. & Kit. [also as R. amoenus Koehler], R. hypomalacus Focke, R. laciniatus Willd., R. lejeunei Weihe & Nees, R. lentiginosus Nees, R. lindbergii Mueller, R. menkei Weihe & Nees, R. mucronulatus Boreau, R. plicatus Weihe & Nees, R. pyramidalis Kaltenb., R. radula Weihe ex Boenn., R. rhombifolius Weihe ex Boenn., R. rudis Weihe & Nees, R. scaber Weihe & Nees [as R. tereticaulis P.J.Mueller], R. serpens Weihe ex Lej., R. silvaticus Weihe & Nees, R. sprengelii Weihe, R. vestitus Weihe & Nees [also as R. conspicuus Mueller], R. villicaulis Koehler ex Weihe & Nees, R. vulgaris Weihe & Nees. Distribution: cosmopolitan; reported in Europe in: Au, Az, Br, Bk, Bu, CS, Ga, Ge, Hb, He, Ho, Hu, It, No, Rm, Su. Common. Leucotelium Tranzschel: Another small genus with only three reported species. According to Cummins & Hiratsuka (1983, 2003) the genus should be included in Sorataea Syd.. However, the genus delimitation here is far from satisfactory, and the author followed an earlier publication by Thirumalachar & Cummins (1948). Cummins & Hiratsuka (1983) state two further species of this genus on Prunus. None of these have so far been reported in Europe. 13. Leucotelium cerasi (Berenger) Tranzschel (1935). SYNONYMS: Mycogone cerasi Berenger (1844); Puccinia cerasi (Berenger) Cast. (1845); Sorataea cerasi (Berenger) Cummins & Y. Hirats. (1983). Gäumann (1959): [799]; Majewski (1977): 286; Braun (1982): 230; Poelt (1985): 62; Minkevicius & Ignataviciute (1991): 169. Spermogonia: epiphyllous; subcuticular; 135 µm diam., conical. Aecia: hypophyllous; distributed over the whole leaf; 0.5 mm; peridium with wide rim; peridial cell walls 7-10 µm thick on outside, smooth; 3-4 µm thick on inside, verrucose. Spores: in chains; 16-26 µm diam., globoid; wall thickness: 1-1.5 µm thicker at apex; finely verrucose; germpores: indistinct. Uredinia: hypophyllous; small; in irregular spots; yellow; aparaphysate. Spores: single; 17-30 × 15-20 µm; globoid to pyriform; wall thickness: 2 µm; yellowish brown; echinulate with irregular glabrous patches; germpores: indistinct. Telia: similar to uredinia. Spores: 2-celled; 30-45 × 15-20 µm; broadened at apex; slightly constricted; wall thickness: 1-2 µm; colourless; smooth; germpores: indistinct; contents: colourless; pedicels: long (to 40 µm); +/- persistent. Life cycle: heteroecious; macrocyclic. Hosts: 0&I: Eranthis Salisb.; II&III: Prunus armeniaca L., P. avium L., P. cerasifera Ehrh., P. cerasus L., P. domestica L., P. dulcis (Miller) D.A.Webb, P. fruticosa Pallas, P. padus L., P. persica (L.) Batsch, P. spinosa L., P. tenella Batsch, P. virginiana L. Distribution: Europe: Au, Bk, Bu, CS, Ge, Hs, Hu, It, Lu, UM; Asia. Uncommon. 341 Ochropsora Dietel: Similar to Tranzschelia, species of this genus produce systemic aecia on Ranunculaceae (but also Fumariaceae) and teliospores on Rosaceae (but also Fabaceae, Elaeagnaceae and Araliaceae). The aecial stage of Ochropsora and Tranzschelia (see below) can be distinguished by the wall thickness of aeciospores which for the former is more or less uniform (around 1 µm), whereas for the latter it is clearly thicker at the distal end (to 3 µm). Uredinia of Ochropsora are characteristically cup shaped on account of the peripheral paraphyses, urediniospores are evenly echinulate, their walls evenly thin (1-1.5 µm). In Tranzschelia the uredinia are not cup shaped and the urediniospores are smooth with thickened walls (5-9 µm) at their apex. The interpretation on a higher taxonomic level has seen some interesting discussions. Gäumann (1959) and Gjærum (1974) record the genus among the Pucciniaceae (with reduced teliospores) whereas most other urediniologists range it with the primitive rusts family Chaconiaceae (Wilson & Henderson 1966, Cummins & Hiratsuka 1983, 2003). 14. Ochropsora ariae (Fuckel) Ramsb. (1914). SYNONYMS: Melampsora ariae Fuckel (1869); Ochropsora sorbi (Oudem.) Dietel. Gäumann (1959): 1216; Wilson & Henderson (1966): 11; Gjærum (1974): 82; Majewski (1977): 155; Braun (1982): 237; Poelt (1985): 63; Minkevicius & Ignataviciute (1991): 75. Spermogonia: epiphyllous subcuticular; 120 µm diam., 60-70 µm high; conical; light brown. Aecia: hypophyllous scattered over the whole surface of the leaves; 0.4 mm; peridium cup shaped, torn revolute; white; peridial cells square; with thick walls: outer 6-10 µm almost smooth; inner 3-5 verrucose. Spores: in chains; 17-26 × 16-21 µm; globoid to ellipsoid; wall thickness: 1 µm; subhyaline; very finely verrucose (>1.2 spines/ µm2); germpores: indistinct; contents: colourless. Uredinia: hypophyllous in chlorotic spots; scattered or in groups; 0.1-0.3 mm diam., surrounded by paraphyses forming a dense circle at the edge of the sorus, these remain when the sorus has shed its spores, forming circular cups; yellowish white; paraphyses broadly cylindrical or clavate 60 µm long 14-18 µm wide. Spores: single; 19-28 × 15-25 µm; ellipsoid or obovoid; wall thickness: 1-1.5 µm; light brown; echinulate (0.77 spines/µm2); germpores: not visible. Telia: hypophyllous initially covered by epidermis; 0.3-0.5 mm; crusty pustules; pale whitish pink. Spores: sessile; at first one-celled later dividing into four cells; 30-70 × 1018 µm; cylindrical; wall thickness: 1 µm; colourless; smooth; germpores: not discernible; contents: opaque grey; granular; teliospores germinating immediately on maturity. Life cycle: heteroecious; macrocyclic. Hosts: 0&I: Anemone L. II&III: Amelanchier ovalis Medicus, Aruncus dioicus (Walter) Fernald, Malus domestica Borkh., M. sylvestris Miller, Prunus avium L., P. padus L., P. tenella Batsch, Pyrus communis L., Sorbus aria (L.) Crantz, S. aucuparia L., S. hybrida L., S. intermedia (Ehrh.) Pers., S. latifolia (Lam.) Pers., S. torminalis (L.) Crantz. Distribution: Europe: Au, Ba, Be, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, He, Ho, Hs, Hu, It, No, Po, Rm, RN, Su, UM; Asia. 342 Phragmidium Link: With 15 species this is the most diverse rust genus on Rosaceae in Europe. It also is the one whose members cause the highest number of diseases (Fig. 2). Wahyuno et al. (2001, 2002) recently compared aecial, uredinial and telial characteristics of Phragmidium species on roses and identified seven distinct aecial types, but found continuous variation in uredinial characters. Similarly, the basic measurements of teliospores did not show discontinuous variation. However, they conclude (Wahyuno et al. 2001) that the putative rusts of ornamental roses do not appear to be a single variable species, and whilst morphological character combinations are not sufficient to differentiate between taxa other properties (such as host plants) may be used to separate them. Many hybrids and subspecific taxa are known in the genera Rubus, Rosa and Potentilla (as well as Alchemilla, see Trachyspora below). The rust infections on these are poorly documented (but see Brandenburger 1994 pp. 31-34 and Scholler 1996 p.170), and have been omitted from this analysis. However, where subspecific taxa and “related species” are listed under specific names in Flora Europaea (Valentine & Charter 1968) they were included in the species analysis under those names. Key to the species: 1 On Sanguisorba........................................................................................... Ph. sanguisorbae On Potentilla.......................................................................................................................... 2 On Rosa................................................................................................................................. 4 On Rubus............................................................................................................................... 8 2 (1) Teliospores mostly 5-6 celled, smooth................................................................ Ph. potentillae Teliospores mainly 4 celled, with delicate warts at the apex...................................................... 3 3 (2) Teliospore pedicels hygroscopic, long............................................................... Ph. andersonii Teliospore pedicels non-hygroscopic, 15-35 µm................................................. Ph. fragariae 4 (1) Teliospores mostly 2, rarely 3 celled.............................................................. Ph. kamtschatkae Teliospores regularly more than 3 celled.................................................................................. 5 5 (4) Teliospores cylindrical - spindle shaped, 8-14, mostly 10-12 celled..................... Ph. fusiforme Teliospores ellipsoid with apical papilla, mostly 7 celled......................................................... 6 6 (5) Apical papilla short (7 - 13 µm)...................................................................... Ph. mucronatum Apical papilla long (15 - 21 µm).............................................................................................. 7 7 (6) Aecia often on stems and fruit, coalescing to 100 mm.........................Ph. rosae-pimpinellifoliae Aecia on leaves, petioles and twigs, max. 2-3 mm......................................... Ph. tuberculatum 8 (1) Teliospores mostly 4 celled................................................................................. Ph. violaceum Teliospores more than 4 celled................................................................................................ 9 9 (8) Teliospores mostly 8-9 (to 14) celled............................................................... Ph. rubi-idaei Teliospores mostly 5-7, not more than 9 celled...................................................................... 10 10 (9) Teliospore width 30-40 µm........................................................................... Ph. candicantium Teliospores more slender (20-30 µm)................................................................................... 11 11(10) Teliospores thick walled (5-7 µm)....................................................................... Ph. bulbosum Teliospores thin walled (3-4 µm).......................................................................................... 12 12(11) Aeciospores echinulate................................................................................... Ph. acuminatum Aeciospores verrucose......................................................................................... Ph. arcticum 343 15. Phragmidium acuminatum (Fr.) Cooke (1871). SYNONYMS: Aregma acuminata Fr. (1815); P. rubi-saxatilis Liro (1908). Gäumann (1959): 1203; Wilson & Henderson (1966): 98; Gjærum (1974): 84; Majewski (1977): 317; Braun (1982): 238; Poelt (1985): 65; Minkevicius & Ignataviciute (1991): 179. Fig. 3E. Spermogonia: not found. Aecia: hypophyllous occasionally also epiphyllous; single or in groups; small (0.30.5 mm diameter); circular caeomata; elongated on the nerves; surrounded by numerous paraphyses (50-75 × 5-8 µm). Spores: 15-29 × 14-26 µm; globoid to ellipsoid; wall thickness: 2-3 µm; yellow; verrucose to echinulate; germpores 3-7. Uredinia: hypophyllous; scattered; small; circular; yellow; similar to aecia. Spores: single; 23-29 × 20-23 µm; globoid; wall thickness: 2-3 µm; colourless; echinulate; germpores: indistinct. Telia: hypophyllous; single or in groups; small; circular at times confluent; black. Spores: 4-8, generally 6-7 celled; 50-110 × 20-34 µm; cylindrical with apical papilla (7-18 µm long); wall thickness: 3-5 µm; brown; densely verrucose; germpores: 2-4 per cell; pedicels: hygroscopic; mostly longer than the spore; persistent. Fig. 3E. Life cycle: Autoecious; macrocyclic. Host: Rubus saxatilis L. Distribution: Europe: Au, Br, Bu, CS, Da, Fe, Ge, Is, No, Rm, RE, UM; northern Asia. Uncommon in Europe. This rust is very similar to P. arcticum and P. bulbosum, the latter has a lower teliospore cell count, also the aeciospores are more clearly echinulate in the present species, compared with variably verrucose aeciospores in P. arcticum or P. bulbosum. 16 Phragmidium andersonii Shear (1902); [Gäumann (1959): 1179]; Gjærum (1974): 85; Minkevicius & Ignataviciute (1991): 188. Spermogonia: not found. Aecia: hypophyllous; 0.5 mm diam; circular caeomata; orange; some paraphyses surrounding aecium. Spores: 21-28 × 15-25 µm; ellipsoid; wall thickness: 1.5-2.5 µm; dark yellow; verrucose. Uredinia: hypophyllous; 0.3 mm; circular; surrounded by many paraphyses. Spores: single; 18-29 × 14-22 µm; obovoid to ellipsoid; wall thickness: 1.5-2.5 µm; dark yellow; finely verrucose. Telia: epi- and hypophyllous; black. Spores: 3-5-celled; 44-85 × 25-35 µm; cylindrical with short apical papilla (3-5 µm); wall thickness: 3-4 µm; dark brown; germpores: 2-3 per cell; pedicels: hygroscopic; long; colourless; persistent. Life cycle: autoecious; macrocyclic. Host: Potentilla fruticosa L. Distribution: Europe: Ba, No, RN, Su; North America; northern Asia. Very rare in Europe. 344 The distribution of Potentilla fruticosa is mainly Asian / North American, with few natural localities in Europe. The rust has only been found in the Baltic (Gäumann 1959), Norway and Northern Russia, but could occur on cultivated P. fruticosa. 17. Phragmidium arcticum Lagerh. (1908); Gäumann (1959): 1205; Gjærum (1974): 86; Minkevicius & Ignataviciute (1991): 178. Spermogonia: not found Aecia: hypophyllous; 0.5 mm diam. circular caeomata; elongated along the veins; surrounded by colourless paraphyses. Spores: 18-24 µm; globoid; wall thickness: 2.5 µm; irregularly verrucose to echinulate; germpores 3-4 indistinct. Uredinia: hypophyllous; small; circular; surrounded by paraphyses. Spores: single; 20-30 × 16-24 µm; obovoid; wall thickness: 1-1.5 µm; colourless; finely echinulate; germpores: 2-3 indistinct. Telia: hypophyllous; black; surrounded by paraphyses similar to the other sori. Spores: 4-9, normally 6-7-celled; 65-100 × 22-30 µm; cylindrical with apical papilla (5-12 µm); wall thickness: 3-4 µm; dark brown; germpores: 2-4 per cell; pedicels: hygroscopic; 130 µm long; colourless; persistent. Life cycle: autoecious; macrocyclic. Host: Rubus arcticus L. Distribution: arctic Europe: Fe, No, RN, Su and Asia. Rare. Closely related to P. acuminatum and P. bulbosum this species occupies a niche on the boreal Rubus arcticus. 18. Phragmidium bulbosum (F.Strauss) Schltdl. (1824). SYNONYMS: Uredo bulbosum F.Strauss (1810); P. rubi (Pers.) Q.Winter (1881). Gäumann (1959): 1200; Wilson & Henderson (1966): 95; Gjærum (1974): 86; Majewski (1977): 314; Braun (1982): 238; Poelt (1985): 65; Minkevicius & Ignataviciute (1991): [182]. IMI 203. Fig. 3B. Spermogonia: epiphyllous; subcuticular; in small groups; 75-100 µm diam; 40 µm high; flat conical; in yellow or reddish spots. Aecia: hypophyllous single or in small groups; 0.5-1 mm diam. elongated at the veins; often forming a ring around the spermogonial position; orange; surrounded by colourless clavate paraphyses (45-75 × 7-12 µm). Spores: in short chains; 20-25 × 15-22 µm; obovoid to ellipsoid; wall thickness: 1-2 µm; colourless; ornamentation: variable warts, rather large (up to 2-7 × 1-2.5 µm); germpores 2-4; contents: yellow. Uredinia: hypophyllous; scattered; 0.3 mm; confluent at times; circular; surrounded by many clavate, curved paraphyses; yellow; often producing yellowish rarely reddish spots on adaxial leaf surface. Spores: single; 20-28 × 14-21 µm; globoid to ellipsoid; wall thickness: 1-2 µm; yellow; echinulate (0.57-0.8 spines/µm2). Fig. 3B; germpores: 2-4. Telia: hypophyllous; scattered; early exposed; 0.5mm; circular; black. Spores: 2-7celled; mostly 5-6-celled; 30-115 × 25-29 µm; cylindrical; not constricted; with hyaline 345 papilla (to 12 µm); wall thickness: 5-7 µm; brown; verrucose (colourless warts); germpores: 2-4 per cell; pedicels: hygroscopic; hyaline up to 140 µm long; persistent. Life cycle: autoecious; macrocyclic. Hosts: Rubus caesius L., R. candicans Weihe ex Reichenb. [also as R. thyrsanthus Focke], R. canescens DC., R. chloocladus W.C.R.Watson [as R. vestii Focke], R. discolor Weihe & Nees, R. egregius Focke, R. fruticosus L. s.l., R. glandulosus Weihe ex Lej. & Court., R. gratus Focke, R. hirtus Waldst. & Kit., R. macrophyllus Weihe & Nees [as R. montanus Libert ex Lej.], R. nessensis W.Hall, R. plicatus Weihe & Nees, R. questieri P.J.Mueller & Lefèvre, R. radula Weihe ex Boenn., R. sprengelii Weihe, R. sulcatus Vest ex Tratt., R. ulmifolius Schott, R. villicaulis Koehler ex Weihe & Nees. Distribution: Europe: Au, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, Kr, No, Po, Rm, RE, Si, Su, UM; Asia; northern Africa. One of the more common bramble (Rubus) rusts similar to P. violaceum but with a higher number of teliospore cells and a longer papilla than the latter. 19. Phragmidium candicantium (Vleugel) Dietel (1927). SYNONYM: P. rubi (Pers.) Q.Winter var. candicantium Vleugel. Gäumann (1959): 1199; [Wilson & Henderson (1966): 100]; Gjærum (1974): 87; Majewski (1977): 314; Poelt (1985): 65. Spermogonia: unknown. Aecia: unknown. Uredinia: hypophyllous; scattered; small; circular; surrounded by club shaped paraphyses. Spores: single; 21-26 × 16-21 µm; obovoid to ellipsoid; wall thickness: 2-3µm; distantly echinulate (0.38-0.46 spines/µm2); germpores: 4. Telia: hypophyllous; scattered at times confluent; small; circular; black. Spores: 58-celled, mostly 6-7-celled; 70-135 × 30-40 µm; cylindrical; with short hyaline papilla; wall thickness: 4.5-6 µm; brown; verrucose (colourless warts); germpores: up to 4; pedicels: hygroscopic; hyaline; long (@140 µm); persistent. Life cycle: not known. Hosts: Rubus canescens DC., Rubus candicans Weihe ex Reichenb. [as R. constrictus PJ Müller & Lefèvre and R. thyrsanthus Focke]. Distribution: Europe: Au, CS, Ge, Hs, Hu, Lu, No, Su, UM. Uncommon. According to Henderson (2000) this rust has not been reported in the British Isles. Together with its hosts it appears to have a more southerly distribution but has been reported from Norway (Jørstad 1953) and Sweden (Farr et al. 2004). 20. Phragmidium fragariae (DC.) Rabenh. (1855). SYNONYMS: Puccinia fragariae DC. (1808); Ph. granulatum Fuckel (1869); Ph. fragariastri (DC.) J.Schröt. (1887). Gäumann (1959): [1179]; Wilson & Henderson (1966): 100; Gjærum (1974): 88; Majewski (1977): 303; Braun (1982): 238; Poelt (1985): 63; Minkevicius & Ignataviciute (1991): [188]. 346 Spermogonia: epiphyllous subcuticular; sizeable 15-20 µm high; shallow cushions; yellow; often surrounding the aecia when these occur adaxially. Aecia: mainly hypophyllous; occasionally epiphyllous or on the petioles; 0.5-2 mm; round to oblong; orange; surrounded by clavate paraphyses 50-70 × 12-16 µm. Spores: 17-28 × 14-24 µm; obovoid, ellipsoid to angular; wall thickness: 2 µm; hyaline; partly finely partly densely verrucose (warts up to 3 µm); germpores indistinct; contents: orange. Uredinia: hypophyllous; scattered; small; round; surrounded by and including numerous capitate paraphyses (80 × 10-20 µm). Spores: single; 18-25 × 16-22 µm; globoid to obovoid; wall thickness: 2 µm; orange; verrucose; similar to aeciospores; germpores: indistinct. Telia: hypophyllous; scattered; small; circular; black; soon naked; pulverulent. Spores: 2-5-celled mostly 4-celled; 40-85 × 22-28 µm; rounded at the apex; slightly constricted; never papillate; wall thickness: 4 µm; pale brown; with some delicate warts around the apex; germpores: usually 3 occasionally 2; pedicels: non-hygroscopic; hyaline 15-35 µm; persistent. Life cycle: autoecious; macrocyclic. Hosts: Potentilla alba L., P. alchimilloides Lapeyr., ., P. argentea L., P. micrantha Ramond ex DC., P. montana Brot., P. patula Waldst. & Kit., P. pyrenaica Ramond ex DC., P. sterilis (L.) Garcke. Distribution: Europe: Au, Br, Bk, Bu, CS, Da, Ga, Ge, Gr, Hb, Hs, Hu, It, Kr, No, Po, Rm, Su, UM and other northern hemisphere. Common. A member of the sub genus Earlea. The teliospores are almost smooth and without papilla; they resemble Frommeella tormentillae (see above) to some extent, but for the number of germpores in the cells. 21. Phragmidium fusiforme J.Schröt. (1871); Gäumann (1959): 1194; Gjærum (1974): 89; Majewski (1977): 305; Braun (1982): 238 Poelt (1985): 64; Spermogonia: epiphyllous subcuticular; occasionally hypophyllous; 100 µm diam 40 µm high; conical. Aecia: on leaves; petioles and fruits; large on fruits; medium sized on leaf veins and leaves; round on leaves and fruits; oblong on leaf veins and petioles; orange; surrounded by clavate to capitate paraphyses 70-90 × 10-20 µm. Spores: in chains with distinct intercalary cells; 18-30 × 15-21 µm; globoid to angular; wall thickness: 1.5-2 µm; hyaline; loosely echinulate - verrucose (0.58 warts/µm2); germpores several, indistinct; contents: orange. Uredinia: hypophyllous; scattered or in groups; small; punctiform; coalescing at times; surrounded by paraphyses which are often curving inwards (50 × 8-11 µm). Spores: single; 17-27 × 15-21 µm; globoid to ellipsoid; wall thickness: 1.5-2 µm; hyaline; echinulate (1.15 spines/µm2); germpores: obscure. Telia: hypophyllous; originating in the uredinia; small; punctiform; coalescing at times; black; as uredinia. Spores: (1-)8-14, mostly 10-12-celled; 42-114 × 21-31 347 µm; cylindrical to spindle shaped; with conical apex; individual cells short; wall thickness: 4-5 µm; brown; with numerous hyaline warts; germpores: 2-4; pedicels: hygroscopic; hyaline; 60-160 µm; broad at the base (15-18 µm); persistent. Life cycle: autoecious; macrocyclic. Hosts: Rosa acicularis Lindley, R. gallica L., R. glauca Pourret, R. glutinosa Sibth. & Sm., R. majalis J. Herrmann, R. mollis Sm., R. pendulina L., R. villosa L. Distribution: Europe: Au, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, He, Hs, Hu, It, No, Po, Rm, UM and other northern hemisphere. Fairly common. The most remarkable rose rust in respect of its teliospore whilst being relatively variable in cell number it exhibits a consistent and unmistakable spindle shape and relatively flat, disk shaped cells. In their addition to the British rust flora Henderson & Bennell (1979) include this species for the British Isles (collected by Plowright 1889, p.226, as P. rosae-alpinae). 22. Phragmidium kamtschatkae (F.W.Anderson) Arthur & Cummins (1933). SYNONYMS: Puccinia kamtschatkae F.W.Anderson (1890) Puccinia rosae Barclay (1889) nom. illegit. Gäumann (1959): [928]; Sydow (1904) 487. Spermogonia: epiphyllous; covering the whole surface of the leaf +/- evenly; producing unpleasant smell whilst active. Aecia: unknown. Uredinia: unknown. Telia: amphigenous; coalescing; soon naked; pulverulent; dark brown; covering leaves and branches; causing affected leaves to thicken; perennial mycelium; causing loss of shoots and eventually whole plants. Spores: 2-celled; at times 3-celled; 30-50 × 16-35 µm; ellipsoid to oblong; apex round not thickened; slightly constricted; base rounded rarely tapering slightly; yellow; verrucose; germpores: 1 per cell; occasionally 2 per cell; pedicels: short; hyaline to yellow; +/- persistent. Life cycle: autoecious; microcyclic. Host: Rosa acicularis Lindley. Distribution: north-eastern Europe: Fe, RN; western Asia. This rust is listed in the IUCN list as critically endangered (Anon 2004b). Gäumann (1959) includes this rust in his ‘Formenkreis’ of Puccinia tartrensis (= P. sieversiae; see below), on account of the many two-celled teliospores of this fungus. It clearly is not a typical Phragmidium. The mycelium of this fungus is perennial, and it can therefore cause considerable dieback in the affected host (Sydow & Sydow 1904). 23. Phragmidium mucronatum (Pers.) Schltdl. (1824). SYNONYMS: Puccinia mucronata a Puccinia rosae Pers. (1801); Ph. disciflorum (Tode)J.James (1895). Gäumann (1959): 1190; Wilson & Henderson (1966): 104; Gjærum (1974): 90; Majewski (1977): 308; Braun (1982): 239; Poelt (1985): 64; Minkevicius & Ignataviciute (1991): [184]. IMI 204. 348 Spermogonia: epiphyllous; subcuticular; small (110-115 µm diam. 35-40 µm high); flat; yellow. Aecia: hypophyllous on leaves; on petioles and fruits; 1mm but often coalescing on branches and than large (to 10mm); circular on leaves; irregular on veins or petioles or branches or fruits; orange; surrounded by clavate to capitate paraphyses 70-80 × 8-18 µm. Spores: 20-30 × 16-21 µm; globoid to ellipsoid; wall thickness: 1.5-3 µm; hyaline; distantly verrucose (0.45-0.58 warts/µm2); germpores 6-8. Uredinia: hypophyllous; scattered or in groups; small (0.1-0.6 mm); circular; soon naked; pale orange; surrounded by paraphyses which are often curving inwards (70 × 7-15 µm). Spores: single; 20-28 × 14-21 µm; globoid to ellipsoid; wall thickness: 1.5-2 µm; hyaline; echinulate (0.77 spines/µm2); germpores: around 8 scattered; 22.5 µm in diam. Telia: hypophyllous; originating in the uredinia; small; punctiform; coalescing at times; black or brown; as uredinia. Spores: 5-9, mostly 6-8 celled; 65-110 × 22-30 µm; ellipsoid to fusoid; hardly constricted; tapering apically with papilla (7-13 µm long); wall thickness: 5-7 µm; dark brown; coarsely verrucose; germpores: 2-3; pedicels: hygroscopic; hyaline; long (130 µm) swollen at base (22-27 µm); persistent. Life cycle: autoecious; macrocyclic. Hosts: Rosa agrestis Savi, R. arvensis Hudson, R. blanda Aiton, R. caesia Sm., R. canina L., R. corymbifera Borkh., R. elliptica Tausch, R. foetida J. Herrmann, R. gallica L., R. glauca Pourret, R. jundzillii Besser, R. majalis J. Herrmann, R. micrantha Borrer ex Sm., R. mollis Sm., R. orientalis Dup., R. pimpinellifolia L., R. rubiginosa L., R. rugosa Thunb., R. sempervirens L., R. subcanina D.H. Christ, R. tomentosa Sm., R. villosa L., R. virginiana J. Herrmann, R. vosagiaca Desportes. Distribution: cosmopolitan in Europe reported from: Au, Az, Bl, Br, Bu, Cr, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, Is, Kr, Lu, No, Po, Rm, RE, Su, UM. Very common. The rose rust. Arguably the most economically important rust on ornamental roses (Shattock 1988). Its control is fairly well documented (Shattock & Rahbar Bhatti 1983). 24. Phragmidium potentillae (Pers.) P.Karst. (1879). SYNONYM: Puccinia potentillae Pers. (1801). Gäumann (1959): 1181; Wilson & Henderson (1966): 101; Gjærum (1974): 91; Majewski (1977): 301; Braun (1982): 239; Poelt (1985): 64; Minkevicius & Ignataviciute (1991): 186. Spermogonia: amphigenous; subcuticular; at times on petioles; 80-200 µm often coalescing; 25-45 µm high; round cushions; yellow; often surrounded by caeomata. Aecia: mostly hypophyllous; also epiphyllous and on the petioles; scattered; 0.5-2 mm forming bigger crusts as they coalesce; +/- circular; orange; surrounded by thin walled clavate to cylindrical paraphyses (80 µm × 6-10 µm). Spores: 18-28 × 14-25 µm; globoid to ellipsoid; wall thickness: 1.5-2 µm; hyaline; finely verrucose (0.58 warts/µm2); germpores indistinct; contents: orange. Uredinia: hypophyllous; scattered or in groups; small (0.5-1 mm); +/- circular or elongated; dark orange; at first covered; then surrounded by paraphyses which are 349 often curving inwards (80 × 10-20 µm). Spores: single; 17-28 × 14-25 µm; globoid to obovoid; wall thickness: 1-2 µm; yellow; finely echinulate (0.46-0.58 spines/ µm2); germpores: indistinct. Telia: hypophyllous partly originating in uredinia; 1mm; circular; soon naked; cushion shaped; black; without particular paraphyses (other than uredinial). Spores: 1-7, mostly 5-6 celled; 32-108 × 20-30 µm; cylindrical to clavoid; rounded or papillate; wall thickness: 3-4 µm to 10 µm at the apex; dark brown; smooth; germpores: 2-3 in the upper part of each cell; pedicels: non-hygroscopic; hyaline; long (60-240 µm); 7-12 µm slightly tapering at the base; persistent. Life cycle: autoecious; macrocyclic. Hosts: Potentilla alchimilloides Lapeyr., P. argentea L., P. argyrophylla Wallich ex Lehm., P. atrosanguinea Loddiges ex D. Don, P. aurea L., P. brauniana Hoppe, P. carniolica A. Kerner, P. caulescens L., P. chrysantha Trev., P. chrysantha Trev., P. cinerea Chaix ex Lehm, P. crantzii (Crantz) G. Beck ex Fritsch, P. detommasii Ten., P. heptaphylla L., P. humifusa Willd, P. inclinata Vill., P. intermedia L., P. leucopolitana Mueller, P. longifolia Willd., P. micrantha Ramond ex DC, P. montana Brot., P. multifida L., P. nivea L., P. norvegica L., P. patula Waldst. & Kit., P. patula Waldst. & Kit., P. pedata Willd., P. pensylvanica L., P. pusilla Host, P. recta L., P. tabernaemontani Ascherson, P. ternata C. Koch, P. thuringiaca Bernh. Distribution: northern hemisphere, in Europe reported in: Au, Ba, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, It, Kr, No, Po, RE, Rm, RN, Su, UM. Uncommon. A member of the smooth, non-hygroscopic subgenus Earlea. In contrast to the other Potentilla rusts this species has a short papilla on its teliospores. 25. Phragmidium rosae-pimpinellifoliae Dietel (1905). SYNONYM: Ph. rosarum f. rosae-pimpinellifoliae Rabenh. (1873). Gäumann (1959): 1186; Wilson & Henderson (1966): 103; Gjærum (1974): 92; Majewski (1977): 311; Braun (1982): 239; Poelt (1985): 65; Minkevicius & Ignataviciute (1991): 185. IMI 205. Spermogonia: mainly on shoots; irregularly scattered; often in groups; 50-120 µm diam. 15-30 µm high; yellow. Aecia: on branches on petioles and veins; on fruits; up to 100 mm along branches; irregular often coalescing; bright orange; at times paraphysate. Spores: 18-27 × 1520 µm; globoid to ellipsoid; wall thickness: 2 µm; hyaline; finely verrucose; germpores 6-8; contents: orange. Uredinia: hypophyllous; scattered; irregular; small (0.1 mm); circular; orange; surrounded by strongly incurved paraphyses (30-50 × 8-12 µm). Spores: 18-25 × 16-20 µm; globoid to ellipsoid; wall thickness: 2-2.5 µm; hyaline; closely echinulate; germpores: indistinct; contents: orange coloured. Telia: inside the uredinia or else hypophyllous; small (0.1 mm); circular; brown; sharing the uredinial paraphyses. Spores: 6-8-celled; 65-115 × 25-34 µm; not 350 constricted; with conical hyaline apical papilla (14-20 µm long); wall thickness: 4-6 µm; opaque; brown; finely verrucose especially apically; germpores: 2-3 per cell; pedicels: hygroscopically swelling in water; firm; hyaline; about the same length as the teliospore; up to 25 µm at the base; persistent. Life cycle: autoecious; macrocyclic. Hosts: Rosa canina L., R. foetida J. Herrmann, R. glauca Pourret, R. majalis J. Herrmann, R. pimpinellifolia L., R. rubiginosa L., R. vosagiaca Desportes. Distribution: northern hemisphere, in Europe reported from: Au, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, Hb, Hs, Hu, Kr, No, Po, Rm, RE, Si, Su, UM. Uncommon. Mainly found on sect. Pimpinellifolia but also reported on sections Canina and Rubiginosa (Gäumann 1959, Brandenburger 1963). This rust can cause dramatic dieback as it girdles even thicker branches with its caeomata. 26. Phragmidium rubi-idaei (DC.) P.Karst. (1879). SYNONYM: Puccinia rubi-idaei DC. (1815). Gäumann (1959): 1205; Wilson & Henderson (1966): 96; Gjærum (1974): 93; Majewski (1977): 312; Braun (1982): 239; Poelt (1985): 65; Minkevicius & Ignataviciute (1991): 180. IMI 207. Spermogonia: epiphyllous; subcuticular; 45-90 µm diam. 20-35 µm high; conical; yellow; surrounded by aecia. Aecia: epiphyllous in groups forming small rings (little over 1 mm); surrounding spermogonia; 0.3-1 mm; at times coalescing; caeomata; circular; orange-yellow; surrounded by hyaline clavate incurved paraphyses (40-70 µm × 14-18 µm). Spores: 16-25 × 14-18 µm; obovoid to ellipsoid; wall thickness: 2-3 µm; pale yellow; very sparsely echinulate (lens-shaped base with fine projection); (0.26-0.33 spines/µm2); germpores obscure. Uredinia: hypophyllous; scattered; very small (0.1-0.3 mm); circular or irregular; pale orange; surrounded by hyaline clavate incurved paraphyses (40-70 µm × 14-24 µm). Spores: single; 15-27 × 14-20 µm; broadly ellipsoid; wall thickness: 1-3 µm; hyaline; sparsely echinulate (0.38 spines/µm2); germpores: obscure; contents: orange. Telia: hypophyllous; in small groups; small (0.3-0.7 mm); circular; black; surrounded by hyaline clavate incurved paraphyses. Spores: 1-14, mostly 7-9celled); 75-144 × 28-35 µm; cylindrical; rounded at base; tapering at the apex; hyaline papilla (up to 15 µm long); wall thickness: 3-6 µm; dark brown; coarsely verrucose; germpores: 3 per cell; pedicels: hygroscopic; longer than spore (to 165 µm); swelling at base (14-27 µm); persistent. Life cycle: autoecious; macrocyclic. Type 11 (Laundon 1972). Hosts: Rubus idaeus L., R. spectabilis Pursh. Distribution: northern hemisphere, in Europe reported from: Au, Ba, Br, Bu, CS, Da, Fe, Ga, Ge, Hb, He, Hs, Hu, It, Lu, No, Po, Rm, RN, Si, Su, UM. Common. In Europe this is the only rust on raspberries (Farr et al. 2004). It can cause economic losses in plantations (Shattock 1988). 351 27. Phragmidium sanguisorbae (DC.) J.Schröt. (1887). SYNONYMS: Puccinia sanguisorbae DC. (1815); Phragmidium poterii Fuckel (1870). Gäumann (1959): [1183]; Wilson & Henderson (1966): 102; Gjærum (1974): 94; Majewski (1977): 321; Braun (1982): 239; Poelt (1985): 64; Minkevicius & Ignataviciute (1991): 186. Spermogonia: amphigenous; subcuticular; circular; rather flat; yellow. Aecia: amphigenous; often forming circles around the spermogonia groups; on veins and petioles; 1mm; longer on veins and petioles; elongated caeomata; orange; surrounded by hyaline clavate incurved paraphyses (40-80 µm × 7-14 µm). Spores: 18-25 × 16-22 µm; globoid to ellipsoid to angular; wall thickness: 1-1.5 µm; hyaline; densely verrucose (0.77-1.15 warts/µm2); germpores 6-8; indistinct; contents: orange. Uredinia: hypophyllous; occasionally epiphyllous; scattered; small (0.25 mm); circular; producing reddish brown spots on adaxial surface; orange; early naked; surrounded by hyaline clavate incurved paraphyses (30-50 µm × 110-170 µm). Spores: single; 17-24 × 15-20 µm; globoid to ellipsoid; wall thickness: 1-1.5 µm; orange; finely echinulate (0.77 spines/µm2); germpores: 6-8 indistinct. Telia: hypophyllous; similar to uredinia; small (0.25-0.5 mm); circular; black; surrounded by hyaline clavate incurved paraphyses (40-60 µm long). Spores: mainly 4-celled; occasionally 2-3- or 5-celled; 40-70 × 20-26 µm; ellipsoid-oblong; slightly constricted at the septa; short apical papilla (5 µm); wall thickness: 3-4 µm; brown; with small scattered hyaline warts; germpores: 2-3 per cell; pedicels: non-hygroscopic; relatively short (21-28 µm long); persistent. Life cycle: autoecious; macrocyclic. Hosts: Sanguisorba minor Scop., S. officinalis L. Distribution: northern hemisphere, in Europe: Au, Bk, Br, Bu, Cr, CS, Da, Ga, Ge, Gr, He, Hb, Hs, Hu, It, Kr, Lu, No, Po, Rm, Si, UM. Fairly common. A member of the subgenus Earlea. Together with Xenodochus carbonarius these are the only rusts on Sanguisorba in Europe. The latter is moderately rare and very easily identified in its teliospore stage (see below). 28. Phragmidium tuberculatum Jul.Müll. (1885); Gäumann (1959): 1188; Wilson & Henderson (1966): 106; Gjærum (1974): 95; Majewski (1977): 307; Braun (1982): 239; Poelt (1985): 65; Minkevicius & Ignataviciute (1991): 183. IMI 208. Spermogonia: epiphyllous; subcuticular; in small groups; 90-100 µm diam. 20-40 µm high; conical; yellow. Aecia: hypophyllous and on leaf veins; on branches and petioles; small on leaves (1 mm); larger on branches and petioles (2-3 mm); circular on leaves; elongated on branches and petioles; yellow; surrounded by hyaline cylindrical incurved paraphyses. Spores: 20-34 × 18-24 µm; globoid to ellipsoid; wall thickness: 1.5-2.5 µm; hyaline; densely verrucose (up to 2 µm wide; average 0.46 warts/µm2); prismatic warts; germpores 6-8 large (4.5 µm diam.); hemispherical swelling inwards. Uredinia: hypophyllous scattered or in groups; small (0.25 mm diam.); circular; pale yellow; surrounded by strongly incurved paraphyses (30-50 × 6-18 µm). Spores: 352 single; 18-30 × 14-23 µm; globoid to ellipsoid; wall thickness: 1.5 µm; hyaline; verrucose - echinulate (0.58 spines/µm2); germpores: 5-8 similar to aecial pores; contents: orange. Telia: hypophyllous; scattered or in groups; minute to larger (to 1.5 mm); at times coalescing; circular to elongated; soon naked; pulverulent; black. Spores: 2-9-celled (mainly 5-7-celled); 40-138 × 22-40 µm; cylindrical to ellipsoid-oblong; hardly constricted at the septa; apical papilla (15-22 µm); wall thickness: 5-7 µm; dark brown; with small scattered hyaline warts; germpores: 2-3 per cell; pedicels: hygroscopic; as long as the spore; hyaline; thickening at the base (to 14-22 µm); persistent. Life cycle: autoecious; macrocyclic. Hosts: Rosa acicularis Lindley, R. agrestis Savi, R. arvensis Hudson, R. caesia Sm., R. canina L., R. chinensis Jacq., R. corymbifera Borkh., R. gallica L., R. glauca Pourret, R. glutinosa Sibth. & Sm., R. majalis J. Herrmann, R. micrantha Borrer ex Sm., R. multiflora Thunb., R. obtusifolia Desv., R. rubiginosa L., R. rugosa Thunb., R. sicula Tratt., R. tomentosa Sm., R. villosa L., R. vosagiaca Desportes.. Distribution: cosmopolitan, in Europe reported from: Au, Az, Ba, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, Kr, Lu, No, Po, Rm, RE, Si, Su, UM. Common. This rust has often been confused with P. mucronatum, the common rose rust. It is not sufficiently differentiated by the number of teliospore cells, however, the abrupt attachment of the teliospore papilla as well as the large size of its urediniospore germ pores make identification possible. 29. Phragmidium violaceum (Schultz) G.Winter (1880). SYNONYM: Puccinia violacea Schultz (1806). Gäumann (1959): 1196; Wilson & Henderson (1966): 98; Gjærum (1974): 96; Majewski (1977): 319; Braun (1982): 240; Poelt (1985): 66; Minkevicius & Ignataviciute (1991): 181. IMI 209. Spermogonia: epiphyllous; subcuticular; densely crowded in dark conspicuous spots; small; hemispherical. Aecia: hypophyllous; scattered or grouped; 1 mm to 10 mm; circular to elongated; orange-yellow; surrounded by clavate +/- straight paraphyses 60 × 18 µm. Spores: in short chains; 19-30 × 17-24 µm; obovoid to ellipsoid; wall thickness: 3-4 µm; hyaline; very sparsely aculeate-verrucose (lens-shaped base with fine projection); (0.23-0.3 warts/µm2); germpores obscure; contents: yellow. Uredinia: hypophyllous; similar to aecia; small; circular or elongated as spots coalesce; orange yellow; surrounded by hyaline clavate or capitate incurved paraphyses (4560 µm × 14-22 µm). Spores: single on pedicels; 20-30 × 17-25 µm; globoid to ellipsoid; wall thickness: 3-4 µm; hyaline; distantly and coarsely aculeate-verrucose (0.23-0.3 spines/µm2); germpores: obscure; contents: yellow. Telia: hypophyllous; small to larger (1.5 mm diam.); circular to elongated, as spots coalesce; soon naked; pulverulent; black. Leaves become purplish red in the area surrounding sori and on the adaxial surface. Spores: 1-5-celled (mostly 4-celled); 55-110 × 30-35 µm; cylindrical to ellipsoid-oblong; hardly constricted at the septa; 353 very short papilla; wall thickness: 6-9 µm; brown; covered in numerous hyaline warts; germpores: 3-4 per cell; pedicels: hygroscopic; hyaline; swollen at base; longer than the spore (up to 180 µm long; 18 µm wide); persistent. Life cycle: autoecious; macrocyclic. Hosts: Rubus albiflorus Boulay & Lecand ex. Coste, R. adscitus Genev. [R. macrothyrsus Lange], R. affinis Weihe & Nees [as R. senticosus Koehler ex Weihe], R. arrhenii Lange, R. bifrons Vest ex Tratt., R. boraeanus Genev. [as R. macrostachys PJ Müller], R. caesius L., R. candicans Weihe ex Reichenb. [also as R. thyrsanthus Focke], R. canescens DC., R. chloocladus W.C.R.Watson [as R. vestii Focke], R. chlorotyrsos Focke [ as R. axillaris Lej., R. cimbricus Focke], R. discolor Weihe & Nees, R. divaricatus P.J.Mueller, R. egregius Focke, R. foliosus Weihe & Nees, R. fruticosus L., R. fuscater Weihe & Nees, R. fuscus Weihe & Nees, R. glandulosus Weihe ex Lej. & Court., R. godronii Lecoq & Lamotte, R. gratus Focke [also as R. sciocharis (Sudre) W.C.R.Watson], R. gremlii Focke [as R. taeniarum Lindeb.], R. hirtus Waldst. & Kit. [also as R. amoenus Koehler], R. hypomalacus Focke, R. infestus Weihe, R. koehleri Weihe & Nees [also as R. bavaricus (Focke) Sudre, R. hebecarpos P.J.Müller], R. laciniatus Willd., R. lejeunei Weihe & Nees, R. lindbergii Mueller [also as R. mercieri Genev.], R. macrophyllus Weihe & Nees, R. mucronulatus Boreau, R. pallidus Weihe & Nees, R. plicatus Weihe & Nees, R. polyanthemus Lindeb., R. pyramidalis Kaltenb., R. radula Weihe ex Boenn., R. rhamnifolius Weihe & Nees [as R. obtusangulus Gremli], R. rudis Weihe & Nees, R. serpens Weihe ex Lej. & Court., R. silvaticus Weihe & Nees, R. sprengelii Weihe [also as R. drejeri Jens.], R. sulcatus Vest ex Tratt., R. ulmifolius Schott [also as R. sanguineus Friv.], R. vestitus Weihe & Nees, R. villicaulis Koehler ex Weihe & Nees [also as R. gelertii Friderichsen, R. septentrionalis W.C.R.Watson], R. vulgaris Weihe & Nees. Distribution: Europe: Al, Au, Az, Be, Bk, Br, Bu, Cr, CS, Da, Ga, Ge, Hb, He, Hs, Hu, It, Kr, Lu, No, Po, Rm, Si, Su, UM; northern Africa. Very common. The common bramble leaf rust - often not recorded because it is thought too ordinary. It mainly attacks the leaves of Rubus sp. in contrast to Kuehneola uredinis which is also found on calyx and stem (see above). Puccinia Pers.: Worldwide the most specious rust genus, Puccinia has two representatives on Rosaceae in Europe. Both are found in Eastern Europe, between Poland / Slovakia and Byelarus / Ukraine. Key to species: Teliospores rugose - pseudoreticulate (Fig. 3H).......................................... Puccinia sieversiae Teliospores smooth.............................................................................. Puccinia waldsteiniana 30 Puccinia sieversiae Arthur (1904) subsp. tatrensis (Urban) Urban (1967). SYNONYM: Puccinia tatrensis Urban. Gäumann (1959): [928]; Majewski (1979): 57; Poelt (1985): 139. Fig. 3H. Spermogonia: lacking. 354 Aecia: lacking. Uredinia: lacking. Telia: mainly hypophyllous; dark brown. Spores: 2-celled; 29-37 × 18-23 µm; ellipsoid to oblong; slightly constricted; wall thickness: 1.5-2.5 µm at sides; 4-6 µm at apex; cinnamon brown; rugose-pseudo-reticulate (Fig. 3H); germpores: 1 per cell; apical in upper cell, near centre in proximal cell; pedicels: hyaline; short; fragile, non-persistent. Fig. 3H. Life cycle: not established. Host: Geum reptans L. Distribution: Tatra Mountains: CS but probably also Po. Rare. A rare fungus of poorly understood biology. In Europe it is restricted to the Tatra Mountains (Urban 1967), although its host is fairly common. The subspecies sieversiae is a more common fungus in North America. 31 Puccinia waldsteiniae M.A.Curtis (1874); Gäumann (1959): 800; Majewski (1979): 58; Sydow & Sydow (1904) 490. Spermogonia: lacking. Aecia: lacking. Uredinia: lacking. Telia: hypophyllous; 1-2 mm; circular to irregular; pale brown. Spores: two-celled; 34-54 × 12-18 µm; subclavate with round to attenuate apex; slightly constricted; wall thickness: 3-5 µm thickened at apex (to 16 µm); brown; smooth; germpores: indiscernible; pedicels: brownish; long (90 µm); persistent. Life cycle: not established. Host: Waldsteinia geoides Willd. Distribution: Galicia, Po, RB, Rm, UM; North America, Asia. Apparently rare. An obscure fungus whose European distribution has only been reported from Galicia (Gäumann 1959), the Ukraine, Rumania (Majewski 1979) and Belarus (Guyot 1965). In North America, Siberia, China and Japan the fungus has been found on other species of Waldsteinia (Majewski 1979, Hiratsuka et al. 1992). I have not been able to see a specimen of this rust from Europe, and the description above is taken from Gäumann (1959), Majewski (1979) and Sydow & Sydow (1904). Pucciniastrum G.H.Otth: There is some debate about the establishment of other genera within Pucciniastrum. Notably, Sato et al. (1993) and Cummins & Hirastuka (2003) recongnise four genera within Pucciniastrum sensu lato: Calyptospora (on Vaccinium), Thekopsora (mainly on Rubiaceae and Ericaceae but also on Rosaceae) and Naohidemyces (on Vaccinium) as well as Pucciniastrum itself (on a wide range of dicotelydons including Rosaceae). However, as the case is not settled yet, I have treated the three following rosaceous rusts within Pucciniastrum sensu lato. 355 Key to the species: Uredinia and telia conspicuous, on Prunus....................................... Pucciniastrum areolatum Telia indistinct, on Agrimonia......................................................... Pucciniastrum agrimoniae Telia flat cushions, on Rubus............................................................... Pucciniastrum arcticum 32. Pucciniastrum agrimoniae (Dietel) Tranzschel (1895). SYNONYM: Thekopsora agrimoniae Dietel (1890). Gäumann (1959): 48 [as P.a. (DC.) Tranzschel]; Wilson & Henderson (1966): 35; Gjærum (1974): 214; Majewski (1977): 80; Braun (1982): 283; Poelt (1985): 53; Minkevicius & Ignataviciute (1991): 38. Spermogonia: unknown. Aecia: unknown. Uredinia: mainly hypophyllous; in groups or covering the whole surface; small (0.20.5 mm); coalescing at times; covered by epidermis and surrounded by a hemispherical peridium with small pore through which the spores get liberated; orange; peridial cells irregularly angular; wall 1.5-2 µm smooth except at opening where they are thicker walled (2.5-5 µm) and echinulate. Spores: single on pedicels; 15-25 × 12-20 µm; obovoid to ellipsoid; wall thickness: 1-1.5 µm; echinulate (0.58 spines/µm2); germpores: indistinct. Telia: hypophyllous; subepidermal; in proximity to uredinia; small; indistinct; irregular; reddish brown; intercellular. Spores: mainly divided into 4 cells by two anticlinal septa but irregular 2-5-celled; 20-25 µm high 15-25 µm diam. irregularly globular; wall thickness: 2 µm; yellowish brown; smooth; germpores: obscure; pedicels: none. Life cycle: unknown. Hosts: Agrimonia eupatoria L., A. pilosa Ledeb., A. repens L. Distribution: cosmopolitan, in Europe reported from: Au, Ba, Be, Bl, Bk, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, Hs, Hu, It, Lu, No, Po, RB, RE, Rm, Su, UM. Fairly common. The life cycle of this rust is not fully understood, but an alternate host would most likely be among the Pinaceae. The teliospore generation is uncommon. 33. Pucciniastrum arcticum Tranzschel (1895). SYNONYM: Uredo arcticus Lagerh. (1889). Gäumann (1959): 49; Gjærum (1974): 215; Majewski (1977): 82. Spermogonia: unknown in Europe. Aecia: unknown in Europe. Uredinia: hypophyllous; often evenly covering the entire surface of the leaf; small (0.10.25 mm); covered by epidermis and surrounded by a hemispherical peridium with small pore through which the spores get liberated; yellowish; peridial cells irregularly angular; wall 1.5-2 µm smooth except at opening where they are thicker walled (2.5-5 µm) and echinulate. Spores: single on pedicels; 16-24 × 12-16 µm; obovoid to ellipsoid; wall thickness: 1-2 µm; hyaline; finely echinulate; germpores: indiscernible. 356 Telia: hypophyllous; subepidermal; in proximity to uredinia; small (0.1-0.3 mm); circular; flat cushions; brown; intercellular. Spores: 18-25 µm diam. globular to angular; brown; smooth; germpores: obscure; pedicels: none. Life cycle: heteroecious; macrocyclic. Hosts: [Picea canadensis (Mill.) B.S.P. (Hirastuka 1958)]; Rubus arcticus L, R. saxatilis L., possibly also on R. chamaemorus L. Distribution: circumpolar, in Europe: Fe, No, RN, Su. This species is described in Gjærum (1974) on R. arcticus in Norway and by others in Russia, Finland and Sweden (Farr et al. 2004) and on R. saxatilis in Russia but it has not yet been found on R. chamaemorus in Europe. 34. Pucciniastrum areolatum (Fr.) G.H.Otth (1863). SYNONYMS: Xyloma areolatum Fr. (1817); Thekopsora areolata (Fr.) Magnus (1875). Gäumann (1959): [53]; Wilson & Henderson (1966): 35; Gjærum (1974): 215; Majewski (1977): 73; Braun (1982): 283; Poelt (1985): 54; Minkevicius & Ignataviciute (1991): [43]. Spermogonia: subcuticular; abaxial on cone scales; also found on young shoots; up to 4 mm diam. irregular flat crusts; whitish; exuding a sugary liquid with strong smell. Aecia: on both sides of the cone scales; crowded; 1-1.25 mm diam. 0.7-1 mm high; honey comb shaped through crowding; peridium firm; brown; peridial cells irregularly polygonal; 22-30 µm long 22-25 µm wide; external walls extremely thick almost completely displacing cell contents; slightly verrucose; internal walls thinner (2.5-3.5 µm) finely verrucose. Spores: in regular chains; 20-28 × 16-22 µm; globoid to angular; wall thickness: 3-6 µm; reddish; densely and pronounced verrucose with one smooth sector where thickness is 3 µm; germpores not observed. Uredinia: hypophyllous; in groups; individual sori small in groups 1-5 mm diam. irregular; bordered by the fine leaf veins; producing spots reddish brown on adaxial surface purplish on abaxial surface. Spores: 15-24 × 10-16 µm; irregular; obovoid to ellipsoid; wall thickness: 1.5-2 µm; hyaline; finely echinulate (0.58 spines/µm2); germpores: indistinct; contents: orange yellow; pedicels: short. Telia: mainly epiphyllous; occasionally hypophyllous; covering the inter-veinal space; sometimes 10 × 10 mm; angular; in crusts; dark brown; glossy; intercellular. Spores: mainly divided into 4 cells by two anticlinal septa but irregular 2-5-celled; 22-30 × 8-14 µm; irregularly globular; divided by anticlinal walls; wall thickness: basal 1 µm; apical 2-3 µm; light brown; smooth; germpores: 1 in the corner where the anticlinal walls meet; pedicels: none. Life cycle: heteroecious; macrocyclic. Hosts: 0&I: Picea spp. II&III: Prunus avium L., P. cerasus L., P. domestica L., P. mahaleb L., P. padus L., P. serotina Ehrh., P. spinosa L., P. virginiana L. Distribution: Europe: Au, Ba, Br, Bu, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hu, It, No, Po, Rm, RN, Su, UM; Asia. Uncommon. 357 Trachyspora Fuckel: A small genus of four species world wide, all of which occur on Alchemilla (Cummins & Hiratsuka 2003). In Europe two species and one variety are recognised. Key to the species: 1 Primary uredinia developing from systemic mycelium, hypophyllous................ T. alchemillae Primary uredinia lacking......................................................................................................... 2 2 (1) Teliospores typically 30+/-2 × 28+/-2 µm............................................................ T. melospora Teliospores typically 35+/-2 × 28+/-2 µm................................. T. melospora var. pentaphyllea 35. Trachyspora alchemillae (Pers.) Fuckel (1861). SYNONYMS: Uredo alchemillae Pers. (1801); Trachyspora intrusa (Grev.) Arthur (1934). Gäumann (1959): 215; Wilson & Henderson (1966): [364]; Gjærum (1974): [223]; Majewski (1977): 267; Braun (1982): 285; Poelt (1985): [142]; Minkevicius & Ignataviciute (1991): 113. Spermogonia: epiphyllous; subepidermal; size not recorded. Uredinia I: hypophyllous; from systemic mycelium; covering whole leaf surface; often coalescing; rounded or elongated; orange; covered by large fragments of the torn epidermis. Spores: single on pedicels; 16-25 × 14-21 µm; globoid to ellipsoid; wall thickness: 1 µm; hyaline; finely echinulate; germpores indistinct; contents: orange; pedicels: short; fragile, non-persistent. Uredinia II: uncertain; if present then as uredinia I. Spores: if present then as primary urediniospores. Telia: hypophyllous; at first originating from aecia; later in separate sori; size as aecia at first; later independent and small; circular; light brown; from same mycelium as aecia later originating from limited independent mycelium. Spores: single celled on short pedicels; 20-40 × 20-30 µm; globoid to obovoid; wall thickness: 3-4 µm; light brown; irregularly and coarsely verrucose on top; lower part smooth to verrucose; germpores: indistinct; pedicels: short (20-40 µm) fragile with single septum; hyaline, non-persistent. Life cycle: autoecious; brachycyclic. Hosts: Alchemilla acutidens Buser, A. acutiloba Opiz, A. alpina L., A. bulgarica Rothm. [also as A. sericata Reichenb.], A. catachnoa Rothm., A. colorata Buser, A. coriacea Buser, A. crinita Buser, A. filicaulis Buser, A. fissa Günther & Schummel, A. flabellata Buser, A. glabra Neygenf., A. glaucescens Wallr., A. glomerulans Buser, A. gracilis Opiz, A. heterophylla Rothm., A. indivisa (Buser) Rothm., A. lapeyrousii Buser, A. monticola Opiz, A. murbeckiana Buser, A. othmarii Buser [as A. gracillima Rothm.], A. pyrenaica Dufour, A. splendens Christ ex Favrat, A. vetteri Buser, A. wichurae (Buser) Stefánsson [also as A.. oxyodonta (Buser) C.G.Westerlund], A. xanthochlora Rothm. Distribution: Northern hemisphere, Europe: Au, Be, Bk, Br, Bu, CS, Da, Fa, Fe, Ga, Ge, Gr, He, Hs, Hu, Is, It, Kr, No, Po, RB, RE, Rm, RN, Su, UM; Southern Africa. Frequent. This rust persists in the rhizomes of its host and leads to early infections in the spring. 358 36. Trachyspora melospora (Therry) Tranzschel (1914). SYNONYM: Uredo melosporus Therry (1873/4). Gäumann (1959): 217; Poelt (1985): 143. Fig. 3G. Spermogonia: not recorded. Uredinia I: lacking; primary urediniospores scattered within telia; 21-28 × 18-21; globoid to ellipsoid; wall thickness: 1 µm; hyaline; finely echinulate; germpores indistinct; contents: orange; pedicels: short; fragile. Uredinia II: as uredinia I. Telia: hypophyllous; small; circular; rusty brown. Spores: single celled on short pedicels; 27-32 × 26-30 µm; globoid to obovoid; wall thickness: 3-4 µm; light brown; irregularly and coarsely verrucose on top; rarely smoother; germpores: indistinct; pedicels: short (20-40 µm) fragile with single septum; hyaline; non-persistent. Fig. 3G. Life cycle: autoecious; microcyclic with some remaining aeciospores. Hosts: Alchemilla alpina L., A. hoppeana (Reichenb.) Dalla Torre, A. indivisa (Buser) Rothm., A. plicatula Gand. Distribution: central Europe: Au, Ge, He. Locally common, possibly overlooked. This rust has never been reported on the Saxatilis series of Alchemilla and seems to be restricted to the series Hoppeana in section Alchemilla and subsection Chirophyllum. Like 35 above the rust persists in the rhizomes of its hosts and leads to early infections in the spring. 36a. Trachyspora melospora var. pentaphylleae (Gäum.) S. Helfer comb. nov.; Gäumann (1959): 218; Poelt (1985): 143. Spermogonia: not recorded. Uredinia I: lacking; primary urediniospores scattered within telia; 21-28 × 18-21; globoid to ellipsoid; wall thickness: 1 µm; hyaline; finely echinulate; germpores indistinct; contents: orange; pedicels: short; fragile. Uredinia II: as uredinia I. Telia: hypophyllous; small; circular; rusty brown. Spores: single celled on short pedicels; 32-39 × 26-30 µm; globoid to obovoid; wall thickness: 3-4 µm; light brown; irregularly and coarsely verrucose on top; rarely smoother; germpores: indistinct; pedicels: short (20-40 µm) fragile with single septum; hyaline, non-persistent. Life cycle: autoecious; microcyclic with some remaining aeciospores. Host: Alchemilla pentaphyllea L. Distribution: central Europe. Uncommon. Gäumann (1943) established this as a separate species on account of its longer teliospores compared with T. melospora. Its host is in the section Pentaphylleae. There is little support for this among uredinologists (e.g. Anon 2004c, Zwetko 2000), 359 as the only morphological difference is in the spore dimensions, the taxon is here reunited with the previous one as a variety. Specimens examined: see Appendix 2. Tranzschelia Arthur: This genus is represented by three species in Europe. Two of these are heteroecious with Prunus as main host and Anemone as alternate host, and one, T. anemones (Pers.) Nannf. is autoecious on Ranunculaceae. The aecial stage produces a perennial mycelium in the Anemone host, where the rust inhibits flower formation. The main differentiating character for this genus on Prunus is in the teliospore basal cells and their ornamentation. The basal cell of T. discolor being oblong elliptic and nearly smooth, whereas that of T. pruni-spinosae is globoid and coarsely verrucose. Whilst the teliospores of the two species are clearly distinct, some of the data concerning uredinial infections may be confused and should be considered with caution. The aecial stage of Ochropsora (see above) and Tranzschelia can be distinguished by the wall thickness of aeciospores which for the former is more or less uniform (around 1 µm), whereas for the latter it is clearly thicker at the distal end (to 3 µm). In Tranzschelia the uredinia are not cup shaped and urediniospores are smooth with thick (5-9 µm) walls at their apex. Uredinia of Ochropsora are characteristically cup shaped on account of the peripheral paraphyses, urediniospores are evenly echinulate and thin walled (1-1.5 µm). Cummins & Hiratsuka (2003) place this genus into the Uropyxidaceae. 37. Tranzschelia discolor (Fuckel) Tranzschel & M.A.Litv. (1939). SYNONYMS: Puccinia discolor Fuckel (1867); T. pruni-spinosae var. discolor (Fuckel) Dunegan (1938). Gäumann (1959): [204]; Wilson & Henderson (1966): 304; Gjærum (1974): 226; Majewski (1977): 290; Poelt (1985): 143. IMI 287. Fig. 3C. Spermogonia: on both sides of leaves; scattered; small; subcuticular; type 7; black. Aecia: hypophyllous; spread evenly over the whole surface of the leaves; 1.5-2.5 mm; circular; peridium with broad revolute margin which is torn; yellowish orange. Spores: 16-24 µm diam. globoid; wall thickness: 1-2 µm; thickened distally (to 3µm); hyaline; finely verrucose; germpores indistinct; contents: yellow. Uredinia: hypophyllous; scattered or crowded and coalescing; small; circular; cinnamon brown; occasionally on adaxial surface. Spores: single soon detached (pulverulent); 20-40 × 10-19 µm; ellipsoid to obovoid; wall thickness: 5-9 µm at apex, 1.5-2 µm elsewhere; yellow to dark brown at apex; smooth at apex; verrucose or echinulate in lower half; germpores: 3 or 4 equatorial; paraphyses present (brown; capitate) size not recorded. Telia: hypophyllous; singly or in groups; 0.25-0.5 mm; circular; blackish brown; soon naked; dusting. Spores: two-celled; 30-45 × 18-25 µm; deeply constricted and easily fracturing; upper cell globoid, verrucose (0.46-0.58 spines/µm2); lower cell oblong-elliptic, almost smooth; wall thickness: 1.5-2 µm; germpores: close to septum; pedicels: short (up to 40 µm); non-persistent. Fig. 3C. Life cycle: heteroecious; macrocyclic. 360 Hosts: 0&I: Anemone sp.; II&III: Prunus armeniaca L., P. avium L., P. cerasifera Ehrh., P. domestica L., P. dulcis (Miller) D.A.Webb, P. mahaleb L., P. persica (L.) Batsch, P. serotina Ehrh., P. spinosa L. Distribution: cosmopolitan, in Europe reported from: Au, Az, Br, Bu, CS, Da, Ga, Ge, Gr, Hb, Ho, Hu, It, No, Si, Tu, UM. Common. 38. Tranzschelia pruni-spinosae (Pers.) Dietel (1922). SYNONYM: Puccinia pruni-spinosae Pers. (1801). Gäumann (1959): 201; Wilson & Henderson (1966): 307; Gjærum (1974): 227; Majewski (1977): 288; Braun (1982): 286; Poelt (1985): 144; Minkevicius & Ignataviciute (1991): 107. IMI 288. Spermogonia: on both sides of leaves; scattered; small; subcuticular; type 7; black. Aecia: hypophyllous; spread evenly over the whole surface of the leaves; 1.5-2.5 mm; circular; peridium with broad revolute margin which is torn; yellowish orange. Spores: 16-24 µm diam. globoid; wall thickness: 1-2 µm; thickened at apex (to 3 µm); hyaline; finely verrucose; germpores indistinct; contents: yellow. Uredinia: hypophyllous; scattered or crowded and coalescing; small; circular; cinnamon brown; occasionally on adaxial surface. Spores: single on pedicels; soon pulverulent; 20-40 × 12-20 µm; ellipsoid; wall thickness: 1-1.5 µm; thickened at apex to 8 µm; yellowish brown; germpores: 3-4 equatorial; other uredinial characters: paraphyses present (brown; capitate) Telia: hypophyllous; singly or in groups; 0.25-0.5 mm; circular; blackish brown; soon naked; dusting. Spores: two-celled; 35-43 × 19-24 µm; deeply constricted and easily fracturing; both cells globoid; coarsely verrucose (0.46-0.58 warts/µm2); wall thickness: 1.5-2 µm; dark brown; germpores: upper cell apical; lower cell basal; pedicels: hyaline; short (up to 40 µm); non-persistent. Life cycle: heteroecious; macrocyclic. Hosts: 0&I: Anemone spp.; II&III: Prunus armeniaca L., P. avium L., P. brigantina Vill., P. cerasifera Ehrh., P. domestica L., P. dulcis (Miller) D.A.Webb, P. fruticosa Pallas, P. mahaleb L., P. persica (L.) Batsch, P. serotina Ehrh., P. spinosa L., P. virginiana L. Distribution: cosmopolitan, in Europe reported from: Al, Au, Az, Ba, Be, Bk, Br, Bu, Cr, CS, Da, Fe, Ga, Ge, Gr, Hb, He, Hs, Hu, It, Kr, Lu, No, Po, RE, Rm, RN, Si, UM. According to Wilson & Henderson (1966) this rust has often been included with T. discolor because the distinction has not been appreciated in the past. The two rusts are indistinguishable in their aecial stage on Anemone in which they can also be confused with the aecia of Ochropsora ariae. Triphragmium Link: globally there are two species in this genus (Lohsomboon 1990), and both are represented here. The main differences between the two species is the aeciospore ornamentation and the differential development of warts on the teliospores, which are smooth in T. filipendulae and coarsely verrucose in T. ulmariae. Both Filipendula rusts cause severe distortions in their host plants, and are therefore very conspicuous. 361 39. Triphragmium filipendulae Pass. (1875); Gäumann (1959): 1212; Wilson & Henderson (1966): 112; Gjærum (1974): 228; Majewski (1977): 329; Braun (1982): 286; Poelt (1985): 145; Minkevicius & Ignataviciute (1991): 191. Spermogonia: hypophyllous or on the petioles; subcuticular flat (type 11); yellow. Uredinia I: amphigenous on petioles and leaf veins; up to 20 mm long; circular to elongated; orange; causing severe distortions of the leaves. Spores: single on pedicels; 21-29 × 15-23 µm; ovoid to pyriform to ellipsoid; wall thickness: 1.5-2 µm; hyaline; echinulate (0.45-0.58 spines/µm2); germpores 2-3 (?) indistinct; contents: orange. Uredinia II: hypophyllous; scattered; 1mm diam. circular; orange. Spores: single on pedicels; 21-34 × 15-21 µm; pyriform to ellipsoid to ovoid; wall thickness: 1.5-2 µm; hyaline; echinulate (0.45-0.58 spines/µm2); germpores: 2-3 (?) indistinct; contents: orange. Telia: hypophyllous; scattered; very small (0.5 mm); circular (punctiform); black; soon naked; pulverulent. Spores: cells in typical triplets, attached at the tip of a triangle; occasionally two or three in rows; 32-50 × 35-38 × 28-31 µm; faintly triangular; flattened; wall thickness: 2-2.5 µm; brown; smooth or with few small warts around the pores; germpores: 1 per cell apically; pedicels: hyaline; delicate; approximately the same length as the spore, deciduous. Life cycle: autoecious, brachycyclic. Host: Filipendula vulgaris Moench. Distribution: Eurasia. In Europe: Au, Ba, Bk, Br, Bu, CS, Da, Fe, Ge, He, Hs, Hu, It, Kr, No, Po, RE, Rm, Su, UM. Uncommon. 40. Triphragmium ulmariae (DC.) Link (1825). SYNONYM: Puccinia ulmariae DC. (1808). Gäumann (1959): 1210; Wilson & Henderson (1966): 112; Gjærum (1974): 229; Majewski (1977): 327; Braun (1982): 286; Poelt (1985): 145; Minkevicius & Ignataviciute (1991): 192. Fig. 3D. Spermogonia: epiphyllous; subcuticular; in circles; inconspicuous (0.35 mm diam.; 60 µm high); circular; yellow-orange. Uredinia I: amphigenous; on petioles and leaf veins; up to 20 mm long; circular to elongated; orange-yellow; causing severe distortions of the leaves; aparaphysate. Spores: single on pedicels; 22-28 × 15-22 µm; obovoid to ellipsoid; wall thickness: 1.5-3 µm; hyaline; distinctly echinulate (0.38-0.47 spines/µm2); germpores indistinct or lacking; contents: orange. Uredinia II: hypophyllous; scattered; 1mm diam. circular; surrounded by paraphyses; yellow brown; paraphyses 30-50 × 8-15 µm. Spores: single on pedicels; 20-30 × 18-25 µm; ellipsoid to obovoid; wall thickness: 1-1.5 µm; hyaline; echinulate (0.45-0.58 spines/µm2); germpores: obscure; contents: orange; comment: very similar to aecia. Telia: hypophyllous; scattered; occasionally in uredinia and rarely aecia; very small (0.4 mm); circular (punctiform); black; soon naked; pulverulent. Spores: brown +/ -elongate; 33-46 × 26-40 × 22-28 µm; heart shaped; faintly triangular; flattened; 362 wall thickness: 1.5-2.5 µm; dark brown; roughly verrucose (hyaline warts 3 × 1µm); germpores: 1 per cell apically; pedicels: hyaline; delicate; approximately the same length as the spore, non-persistent. Fig. 3D. Life cycle: autoecious, brachycyclic. Host: Filipendula ulmaria (L.) Maxim. Distribution: northern hemisphere, in Europe reported from: Au, Ba, Br, Bu, CS, Da, Fe, Ga, Ge, Hb, He, Hs, Hu, Is, Lu, No, Po, RE, Rm, RN, Su, UM. Common. Xenodochus Schltdl.: Two species are known of this genus, with one representative in Europe. This rust is clearly similar to Phragmidium, and has in fact been included in this genus before (Winter 1884). However, it differs remarkably from all other rosaceous rusts, and a separate genus seems justified. In the older literature the rust is reported to lack the urediniospore stage, however, Sato & Sato (1980) proved experimentally that the fungus possessed caeomoid uredinia. According to these workers the presence of marginal paraphyses depends on the maturity of the sori observed. 41. Xenodochus carbonarius Schltdl. (1826); Gäumann (1959): 1207; Wilson & Henderson (1966): 109 (as Xenodocus carbonarius); Gjærum (1974): 264; Majewski (1977): 322; Braun (1982): 300; Poelt (1985): 166; Minkevicius & Ignataviciute (1991): 190. Spermogonia: subcuticular (intra-epidermal type 10); small; type 10; frequently not observed. Aecia: mainly hypophyllous on leaf veins; occasionally epiphyllous on petioles; scattered; up to 10mm long; caeomata with marginal paraphyses; on yellow or purple spots; paraphyses clavate; yellowish; 30-62 × 8-16 µm. Spores: 17-29 × 13-24 µm; globoid to obovoid; wall thickness: 1.5-2 µm; hyaline; with broad (1 µm) verrucae (1.42 warts/µm2); germpores indistinct or lacking; contents: orange. Uredinia: hypophyllous; small; caeomoid with surrounding paraphyses; on yellow or purple spots; paraphyses clavate; yellowish; 30-62 × 8-16 µm. Spores: 17-29 × 13-24 µm; globoid to obovoid; wall thickness: 1.5-2 µm; hyaline; with broad (up to 1 µm), dense verrucae (1.42 warts/µm 2); germpores: indistinct or lacking; contents: orange. Telia: amphigenous; often adjacent to aecia and uredinia; coalescing to 2-4 mm diam. compound sori; strongly convex; black; soon naked; pulvinate. Spores: 3-22celled; up to 300 µm × 23-30 µm; elongate cylindrical; mostly curved; rounded at top and base; wall thickness: 1.5-3 µm; upper cells dark brown; lower cells lighter brown; smooth; end cell loosely verrucose; germpores: 2 opposing at distal ends; end cell with single apical pore often with a short hyaline papilla; pedicels: short; fairly tough; persistent. Life cycle: autoecious; macrocyclic. Hosts: Sanguisorba minor Scop., S. officinalis L. Distribution: northern hemisphere; in Europe reported from: Au, Be, Br, Bu, CS, Ge, He, Hs, Hu, No, Po, Rm, RN, UM. Uncommon. 363 Excluded: Uromyces antipae Săvul. (1938); Gäumann (1959): 400. Spermogonia: not recorded. Aecia: not recorded. Uredinia: hypophyllous; scattered; yellowish brown; at times in crowded groups. Spores: single on pedicels; 16-27 × 13-16 µm; globoid to ellipsoid; light brown; densely and finely echinulate; germpores: 4-6 equatorial. Telia: hypophyllous; scattered; size various; irregular; blackish brown; naked; pulverulent. Spores: single celled; 23-34 × 16-23 µm; very variable; wall thickness: 2-2.5 µm; brown; apical papilla hyaline; germpores: indiscernible; pedicels: 3-10 µm long; +/- tough. Life cycle: not established. Host: Rosa foetida J. Herrmann. Distribution: Rm. Probably very rare. There is very little information on this rust, and I have not been able to confirm the above data. It has not been included in the formal account as it’s only known host, Rosa foetida is not native to Europe. Uromyces urediniformis (Müll. Hal.) Dietel (1912); Gäumann (1959): 401; Majewski (1977): 190; [Braun (1982): 290]. SYNONYM: Chrysomyxa urediniformis Müll.Hall. Spermogonia: not recorded. Aecia: not recorded. Uredinia: hypophyllous; long covered by epidermis; rather large; aparaphysate; intensely orange; inducing reddish brown spots on adaxial surface. Spores: single on pedicels; 21-27 × 17-21 µm; globoid to pyriform to polyedric; golden yellow. Telia: inside the uredinia. Spores: amongst the urediniopores; 32 × 16-19 µm; hyaline; with apical papilla 2.4 µm diam. germpores: apically. Life cycle: unknown. Hosts: Rubus plicatus Weihe & Nees. Distribution: north central Europe. Apparently rare. The only record for this is in Gäumann (1959) who reports it from northern Germany. Braun (1982) considers it as a dubious species. I have not been able to confirm these records. Conclusions The detailed knowledge of host - parasite distribution is essential to allow estimates of host specificity, epidemic potential of the parasites, and frequency status of the 364 organisms (Helfer 1993). Continued investigation into the presence or absence of parasitic fungi in any host taxa is necessary to establish the presence of possible compatibility or resistance mechanisms; close co-operation between fungal and phanerogam taxonomists is therefore desirable. Whilst all areas of Europe would benefit from further research, the following areas should be prioritised to complete the picture of European rust fungi on Rosaceae: Al, Be, Bl, Co, Cr, Ho, Sa, Sb, Si and Tu. Acknowledgements Thanks are expressed to the curators of BERN, BPI, BRA, K, PRC, PUR, Z, and ZT for the loan of specimens and access to specimen data. The suggestions and corrections presented by two anonymous referees were much appreciated. 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Received 1 March 2004, accepted in revised form 25 February 2005. 367 Appendix 1 Host plants of European rosaceous rusts (rust taxa in brackets): Agrimonia eupatoria (32) Agrimonia pilosa (32) Agrimonia repens (32) Alchemilla acutidens (35) Alchemilla acutiloba (35) Alchemilla alpina (35, 36) Alchemilla bulgarica (35) Alchemilla catachnoa (35) Alchemilla colorata (35) Alchemilla coriacea (35) Alchemilla crinita (35) Alchemilla filicaulis (35) Alchemilla fissa (35) Alchemilla flabellata (35) Alchemilla glabra (35) Alchemilla glaucescens (35) Alchemilla glomerulans (35) Alchemilla gracilis (35) Alchemilla heterophylla (35) Alchemilla hoppeana (36) Alchemilla indivisa (35, 36) Alchemilla lapeyrousii (35) Alchemilla monticola (35) Alchemilla murbeckiana (35) Alchemilla othmarii (35) Alchemilla pentaphyllea (36a) Alchemilla plicatula (36) Alchemilla pyrenaica (35) Alchemilla splendens (35) Alchemilla vetteri (35) Alchemilla wichurae (35) Alchemilla xanthochlora (35) Amelanchier ovalis (3, 4, 8, 14) Aruncus dioicus (14) Cotoneaster integerrimus (5, 7) Cotoneaster nebrodensis (5, 7) Cotoneaster niger (5) Cotoneaster nummularia (5) Crataegus azarolus (4, 5) Crataegus calycina (4) Crataegus laciniata (4, 5) Crataegus laevigata (4, 5) Crataegus monogyna (4, 5, 8) Crataegus nigra (4) Crataegus pentagyna (4) Crataegus sanguinea (4, 5) Cydonia oblonga (4, 5, 8, 11) Filipendula ulmaria (40) Filipendula vulgaris (39) Geum reptans (30) Malus domestica (4, 11, 14) Malus sylvestris (11, 14) Mespilus germanica (5) Potentilla alba (20) Potentilla alchimilloides (20, 24) Potentilla anglica (1) Potentilla argentea (20, 24) Potentilla argyrophylla (24) Potentilla atrosanguinea (24) Potentilla aurea (24) Potentilla brauniana (24) Potentilla carniolica (24) Potentilla caulescens (24) Potentilla chrysantha (24) Potentilla cinerea (24) Potentilla collina (24) Potentilla crantzii (24) Potentilla detommasii (24) Potentilla erecta (1) Potentilla fruticosa (16) Potentilla heptaphylla (24) Potentilla humifusa (24) Potentilla inclinata (24) Potentilla intermedia (24) Potentilla longifolia (24) Potentilla micrantha (20, 24) Potentilla montana (20, 24) Potentilla multifida (24) Potentilla neglecta (24) Potentilla nivea (24) Potentilla norvegica (24) Potentilla patula (20, 24) Potentilla pedata (24) Potentilla pensylvanica (24) Potentilla pusilla (24) Potentilla pyrenaica (20, 24) Potentilla recta (1, 24) Potentilla reptans (1, 24) Potentilla sterilis (20, 24) Potentilla tabernaemontani (24) Potentilla ternata (24) Potentilla thuringiaca (24) Prunus armeniaca (13, 37, 38) Prunus avium (13, 14, 34, 37, 38) Prunus brigantina (38) Prunus cerasifera (13, 37, 38) Prunus cerasus (13, 34) Prunus domestica (13, 34, 37, 38) Prunus dulcis (13, 37, 38) Prunus fruticosa (13, 38) Prunus mahaleb (34, 37, 38) Prunus padus (13, 14, 34) Prunus persica (13, 37, 38) Prunus serotina (34, 37, 38) Prunus spinosa (13, 34, 37, 38) Prunus tenella (13, 14) 368 Prunus virginiana (13, 34, 38) Pyracantha coccinea (3, 5) Pyrus amygdaliformis (9) Pyrus communis (4, 5, 9, 14) Pyrus elaeagrifolia (9) Pyrus nivalis (9) Pyrus pyraster (9) Pyrus salicifolia (9) Pyrus syriaca (9) Rosa acicularis (21, 22, 28) Rosa agrestis (23, 28) Rosa arvensis (23, 28) Rosa blanda (23) Rosa caesia (23, 28) Rosa canina (23, 25, 28) Rosa chinensis (28) Rosa corymbifera (23, 28) Rosa elliptica (23) Rosa foetida (23, 25) Rosa gallica (21, 23, 28) Rosa glauca (23, 25, 28) Rosa glutinosa (21, 28) Rosa jundzillii (23) Rosa majalis (21, 23, 25, 28) Rosa micrantha (23, 28) Rosa mollis (21, 23) Rosa multiflora (28) Rosa obtusifolia (28) Rosa orientalis (23) Rosa pendulina (21) Rosa pimpinellifolia (23, 25) Rosa rubiginosa (25, 28) Rosa rugosa (23, 28) Rosa sempervirens (23) Rosa sicula (28) Rosa subcanina (23) Rosa tomentosa (23, 28) Rosa villosa (21, 23, 28) Rosa virginiana (23) Rosa vosagiaca (23, 25, 28) Rubus adscitus (29) Rubus affinis (12, 29) Rubus albiflorus (29) Rubus arcticus (2, 17, 33) Rubus arrhenii (12, 29) Rubus bifrons (29) Rubus boraeanus (29) Rubus caesius (12, 18, 29) Rubus candicans (12, 18, 19, 29) Rubus canescens (12, 18, 19, 29) Rubus chamaemorus (33) Rubus chloocladus (12, 18, 29) Rubus chlorotyrsos (29) Rubus discolor (18, 29) Rubus divaricatus (12, 29) Rubus egregius (18, 29) Rubus foliosus (29) Rubus fruticosus (12, 18, 29) Rubus fuscater (12, 29) Rubus fuscus (12, 29) Rubus glandulosus (12, 18, 29) Rubus godronii (29) Rubus gratus (12, 18, 29) Rubus gremlii (12, 29) Rubus hirtus (12, 18, 29) Rubus hypomalacus (12, 29) Rubus idaeus (26) Rubus infestus (29) Rubus koehleri (29) Rubus laciniatus (12, 29) Rubus lejeunei (12, 29) Rubus lentiginosus (12) Rubus lindbergii (12, 29) Rubus macrophyllus (18, 29) Rubus menkei (12) Rubus mucronulatus (12, 29) Rubus nessensis (18) Rubus pallidus (29) Rubus plicatus (12, 18, 29) Rubus polyanthemus (29) Rubus pyramidalis (12, 29) Rubus questieri (18) Rubus radula (12, 18, 29) Rubus rhamnifolius (29) Rubus rhombifolius (12) Rubus rudis (12, 29) Rubus saxatilis (2, 15, 33) Rubus scaber (12) Rubus serpens (2, 12, 29) Rubus silvaticus (12, 29) Rubus spectabilis (26) Rubus sprengelii (12, 18, 29) Rubus sulcatus (18, 29) Rubus ulmifolius (18, 29) Rubus vestitus (12, 29) Rubus villicaulis (12, 18, 29) Rubus vulgaris (12, 29) Sanguisorba minor (27, 41) Sanguisorba officinalis (27, 41) Sorbus aria (4, 11, 14) Sorbus aucuparia (4, 5, 6, 10, 11, 14) Sorbus chamaemespilus (6, 10, 11) Sorbus domestica (6, 10) Sorbus hybrida (6, 10, 14) Sorbus intermedia (6, 14) Sorbus latifolia (4, 5, 6, 10, 11, 14) Sorbus meinichii (4) Sorbus minima (4) Sorbus mougeotii (6, 10) Sorbus neglecta (4) Sorbus torminalis (4, 5, 10, 11, 14) Waldsteinia geoides (31) 369 Appendix 2 Trachyspora melospora var. pentaphylleae specimens examined: (Z): Uromyces melosporus (Therry) Sydow, sur Alchemilla pentaphylla, Pâturages rocheux, passage du Gotthard, Canton du Tessin, 16.8.1931, leg. Dr. Eug.Mayor. (ZT): Uromyces melosporus (Th.) Syd., on Alchemilla pentaphyllea, Aletschwald, 8.34. Gm. Trachyspora pentaphyllea Gm. on Alchemilla pentaphyllea, Bettmeralp, August 1937. leg. Tino Gäumann. Trachyspora pentaphyllea Gäumann auf Alchemilla pentaphyllea L., Binntalhütte, 15. Aug. 1971. leg. S.Blumer. Trachyspora pentaphyllea Gäum. auf Alchemilla pentaphylla, Kt. Wallis, Schneetälchen am Grat südwestl. Moosfluh, 10.9.1962. leg. E.Müller. Trachyspora pentaphyllea Gm. auf Alchemilla pentaphyllae, Kt. Graubünden San Bernardino, 25.7.1960. leg. F.Hennen. Trachyspora pentaphyllea Gäumann sur Alchemilla pentaphyllea L., Plattje sur Saas Fee.- Vallée de Saas, Canton du Valais, 15. Aug. 1945. leg. Dr. Eug.Mayor. Trachyspora pentaphyllea Gäumann auf Alchemilla pentaphylla, Kt.Wallis, Aletschreservat Moränenweg, 3.8.1964. leg. E.Müller. Uromyces melosporus (Therry) Sydow sur Alchemilla pentaphyllea L., Pente de l’Eggishorn, Canton du Valais, 11.8.1920, leg. Dr. Eug.Mayor. Trachyspora melospora var. melospora specimens examined: (Z): Sydow, Mycotheca germanica 456 on Alchemilla hoppeana. Uromyces melosporus (Therry) Sydow on Alchemilla hoppeana, Oberhalbstein, 2.7.1920, leg. Hans Schulz?. (ZT): Uromyces melosporus Lagh. on Alchemilla hoppeana, Calfeisental, 19.7.1940, leg. Gäumann. ex Vergleichsherbar J. Poelt Nr. 3609 on Alchemilla hoppeana. Uromyces melosporus Lagh. on Alchemilla hoppeana, Wallis: Lötschental, 13.6.1936, leg. Walo Koch. Uromyces melosporus Lagh. on Alchemilla hoppeana, Grat des Chasseral, 29.7.1937, leg. Gäumann. Trachyspora melospora (Therry) Tranzschel, on Alchemilla hoppeana, Gonzen Gipfel, 29.9.1946, leg. E Sulger. Trachyspora melospora (Therry) Sydow, on Alchemilla hoppeana, Valais, 11.10.1953, leg. Ch. Terrier. Trachyspora melospora Th., on Alchemilla hoppeana, Graubünden, 1.11.1943, leg. Walo Koch (43/8174). Trachyspora melospora Therry, on Alchemilla hoppeana, Graubünden, 24.7.1943, leg. O.Jaag. Uromyces melosporus Sydow, on Alchemilla hoppeana, Midwalden, 8.7.1934, leg. Gäumann & Koch. Trachyspora melospora Therry, on Alchemilla hoppeana, ob Seealpsee, Säntis, 9.7.1943, Leg. O.Jaag. Uromyces melosporus (Therry) Sydow on Alchemilla hoppeana, Bois de Laitmaire, 26.9.1918, leg. E.Mayor. 370

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