Skip to main content

Advertisement

Cercosporoid fungi (Mycosphaerellaceae) 4. Species on dicots (Acanthaceae to Amaranthaceae)

Article metrics

  • 97 Accesses

  • 7 Citations

Abstract

The present paper continues a series of comprehensive taxonomic treatments of cercosporoid fungi (formerly Cercospora s. lat.), belonging to the Mycosphaerellaceae (Ascomycota). The fourth contribution of this series initiates treatments of cercosporoid fungi on dicots and comprises species occurring on hosts belonging the the families Acanthaceae, Actinidiaceae, Adoxaceae, Aizoaceae, Altingiaceae, and Amaranthaceae. The species are described and illustrated in alphabetical order under the particular cercosporoid genera, supplemented by keys to the species concerned. A detailed introduction, a survey of currently recognised cercosporoid genera, a key to the genera concerned, and a discussion of taxonomically relevant characters were published in the first part of this series. The following taxonomic novelties are introduced: Cercospora blepharidicola nom. nov., C. celosiigena sp. nov., C. justiciae-adhatodae sp. nov., C. justiciigena nom. nov., C. sambucicola nom. nov., C. thunbergiigena nom. nov., Cercosporella pseudachyranthis comb. nov., Pseudocercospora cyathulae comb. nov., P. depazeoides comb. nov., P. varia var. viburni-sargentii var. nov., P. viburnicola sp. nov., P. viburni-erosi sp. nov., and P. viburni-nudi sp. nov.

Introduction

True cercosporoid fungi belong to Mycosphaerellaceae (Capnodiales, Ascomycota) and comprise a very large group of plant pathogenic, leaf-spotting, economically relevant species that cause diseases on a wide range of hosts, including numerous cultivated plants. In spite of the enormous relevance of this fungal group, there is no modern comprehensive treatment of Cercospora and allied genera, and the only monograph published by Chupp (1954) is seriously outdated. Therefore, a monographic series with treatments of cercosporoid fungi based on host families was initiated (Braun et al. 2013) with the aim of working towards a comprehensive monograph of this generic complex. So far three contributions have been published: part one dealing with cercosporoid fungi on other fungi (mycophylic taxa), on ferns as well as gymnosperms (Braun et al. 2013); part two dedicated to species on monocots, excluding true grasses (Braun et al. 2014); and part three with a treatment of cercosporoids on hosts of Poaceae (Braun et al. 2015). General chapters with generic descriptions and keys to accepted genera are included in the first part. The present contribution is the first part devoted to cercosporoid fungi on dicots, encompassing species on hosts of the families Acanthaceae, Actinidiaceae, Adoxaceae, Aizoaceae, Altingiaceae, and Amaranthaceae. The structure of part 4 follows the principles circumscribed in part 1 (Braun et al. 2013).

Materials and Methods

The present work is a compilation based on our papers and unpublished data, as well as the global literature. Details of methods are given in the papers cited under references. As far as new examinations are concerned, fungal structures have been examined by standard methods of light microscopy, using an Olympus BX50 microscope, with distilled water and lactic acid as media, but without any staining. If possible, measurements of 30 conidia and other structures have been made at a magnification of ×1000. All illustrations have been prepared by UB. The following abbreviations are used: author names follow Brummit & Powell (1992), journals Bridson (2004a, b), and exsiccatae https://doi.org/www.botanischestaatssammlung.de/DatabaseClient/IndExs/index.jsp (IndExs — Index of Exsiccatae). Taxonomy and nomenclature of plant families, genera and species are based on the “Angiosperm Phylogeny Website” (https://doi.org/www.mobot.org/mobot/research/apweb/), Tropicos database (https://doi.org/www.tropicos.org/), and The Plant List (https://doi.org/www.theplantlist.org).

Taxonomic Treatment

Cercosporoid species on dicots s. lat. ( Acanthaceae to Amaranthaceae )

Acanthaceae

Cercospora

Key to Cercospora species on Acanthaceae
  1. 1

    Conidia in chains, 8–40 × 2–3 µm, (0−)1–3(− 4)-septate, hyaline; on Justicia adhatoda [Adhatoda vasica] ………. Cercospora justiciae-adhatodae

    Conidia formed singly ………. 2

  2. 2(1)

    Stromata large, 20–85 µm; conidiophores short, 10–40 × 3–5 µm, 0–4-septate; conidia short, acicular-subcylindrical, 20–50 × 2–4 µm, 0–5-septate ………. C. balaghatensis

    Stromata lacking or smaller, 10–30 µm diam; and/or conidiophores much longer, 10–500 µm, pluriseptate throughout; conidia longer, 15–360 µm, pluriseptate ………. 3

  3. 3(2)

    Conidia obclavate-cylindrical with obconically truncate base ………. 4

    Conidia consistently acicular, base truncate, or at least longer conidia acicular, mixed with shorter obclavate-cylindrical conidia ………. 6

  4. 4(3)

    Conidiophores short, 10–40 × 2.5–5 µm, 0–1-septate; conidia 25–80 µm long; on Justicia spicigera, Central America (Guatemala) ………. C. jacobiniicola

    Conidiophores longer, about 30–185 µm, with more than two septa; conidia longer, about 20–170 µm ………. 5

  5. 5(4)

    Stromata 30–50 µm diam; conidiophores long and aseptate, 35–120 × 3–6 µm; on Justicia betonica, Asia (India) ………. C. justiciigena

    Stromata lacking or very small, < 25 µm diam; length of the conidiophores similar, but 1–6-septate; on Lepidagathis spp ………. C. lepidagathidis

  6. 6(3)

    Conidia acicular to obclavate-cylindrical, base truncate to obconically truncate ………. 7

    Conidia consistently acicular, base truncate ………. 10

  7. 7(6)

    Conidiophores relatively short, 10–30(− 70) µm; stromata 10–70 µm diam; on Acanthus spp ………. C. acanthi

    Conidiophores longer, 25–155 µm; stromata lacking or small, 10–30 µm; on other hosts ………. 8

  8. 8(7)

    Conidia narrow, 30–150 × 2–4 µm, average < 3 µm wide; on Andrographis spp ………. C. andrographidis

    Conidia wider, 3–5 µm, average > 3 µm; on other hosts ………. 9

  9. 9(8)

    Conidiophores to 195 µm long; on Justicia spp ………. C. justiciicola

    Conidiophores much shorter, to 62.5 µm; on Crossandra spp ………. C. crossandrae

  10. 10(6)

    Stromata well-developed, 10–70 µm diam; conidiophores short, 10–40(− 7) pm; on Acanthus sp ………. C. acanthi

    Stromata lacking or small, about 10–45 µm diam; and/or conidiophores much longer, at least partly longer than 50 µm; on other hosts ………. 11

  11. 11(10)

    Conidiophores 40–310 × 4–8.5 µm; conidia rather broad, 40–360 × 3–8 µm; on Thunbergia spp. ………. C. thunbergiana

    Conidiophores and conidia narrower; conidia about 1.5–5 µm wide; on other hosts or if on Thubergia conidia only 2–3 µm wide ………. 12

  12. 12(11)

    Conidia (2−)2.5–5.5(− 6) µm wide, average > 3 ………. 13

    Conidia narrower, 1.5–4 µm, average < 3 µm ………. 17

  13. 13(12)

    Conidiophores long, 30–500 × 3–8 µm; on Hygrophila spp ………. C. hygrophilae

    Conidiophores much shorter, to 250 µm; on other hosts ………. 14

  14. 14(13)

    Stromata well-developed, about 15–55 µm diam; on Andrographis spp ………. C. andrographidicola

    Stromata lacking or small, about 10–25 µm diam; on other hosts ………. 15

  15. 15(14)

    Lesions formed as brown leaf spots with indefinite margin; on Asystasia spp ………. C. asystasiana

    Leasion visible as greyish white leaf spots; on other hosts ………. 16

  16. 16(15)

    Conidiogenous loci 2–3 µm wide; on Justicia adhatoda ………. C. adhatodae

    Conidiogenous loci somewhat narrower, 1.5–2.5 µm wide; on Strobilathes ………. C. strobilanthis

  17. 17(12)

    Stromata well-developed, to 60 µm diam; on Barleria and Blepharis spp ………. 18

    Stromata lacking or small, 10–25 µm diam; on other hosts ………. 19

  18. 18(17)

    Conidiophores long, 10–210 µm; on Barleria spp ………. C. barleriicola

    Conidiophores much shorter, 15–65 µm; on Blepharis spp ………. C. blepharidicola

  19. 19(17)

    Conidia very narrow, about 1.5–2.5 µm; on Ruellia spp ………. C. ruellina

    Conidia wider, 2–4 µm; on other hosts ………. 20

  20. 20(19)

    Conidiophores 10–100 µm long; on Justicia and Rhytoglossa spp ………. C. diantherae

    Conidiophores longer, to 220 µm; on other hosts ………. 21

  21. 21(20)

    Leaf spots yellowish brown to dark brown, later greyish white with dark border, 0.5–4 mm diam; on Peristrophe spp ………. C. peristrophes

    Leaf spots dark brown to black, vein-limited, 3–10 mm diam; on Thunbergia ………. C. thunbergiigena

Tabular key to Cercospora species on Acanthaceae according to host genera
  • Acanthus

    A single species ………. C. acanthi

  • Adhatoda, see Justicia

  • Andrographis

    1 Conidia acicular, base truncate, 200–240 × 3.3–5 µm ………. C. andrographidicola

    Conidia acicular to obclavate, base truncate to obconically truncate, shorter and narrower, 30–150 × 2–4 µm ………. C. andrographidis

  • Asystasia

    A single species ………. C. asystasiana

  • Barleria

    A single species ………. C. barleriicola

  • Blepharis

    A single species ………. C. blepharidicola

  • Crossandra

    A single species ………. C. crossandrae

  • Hygrophila

    A single species ………. C. hygrophilae

Justicia
  1. 1

    Conidia in chains, 8–40 × 2–3 µm, (0−)1–3(− 4)-septate, hyaline; on Justicia adhatoda [Adhatoda vasica] ………. Cercospora justiciae-adhatodae

    Conidia formed singly, much longer and pluriseptate ………. 2

  2. 2(1)

    Stromata large, 20–85 µm diam; conidiophores short, 10–40 × 3–5 µm, 0–4-septate; conidia short, acicular-subcylindrical, 20–50 × 2–4 µm, 0–5-septate ………. C. balaghatensis

    Stromata lacking or smaller, 10–30 µm diam; conidiophores much longer, 10–195 µm, pluriseptate throughout; conidia longer, 15–250 µm, pluriseptate ………. 3

  3. 3(2)

    Conidia consistently acicular, base truncate, or at least longer conidia acicular, mixed with shorter obclavate-cylindrical conidia ……….. 4

    Conidia obclavate-cylindrical with obconically truncate base, acicular conidia lacking ……….. 6

  4. 4(3)

    Conidia acicular to obclavate-cylindrical, base truncate to obconically truncate ……….. C. justiciicola

    Conidia consistently acicular, base truncate ……….. 5

  5. 5(4)

    Conidia, 35–250 × 2.5–5 µm; on Justicia adhatodae [Adhatoda vasica], Asia (India) ……….. C. adhatodae

    Conidia narrower, 40–180 × 2–4 µm; on Justicia spp., common in North America (records from Africa and Asia rare and unproven) ……….. C. diantherae

  6. 6(3)

    Stromata 30–50 µm diam; conidiophores long and aseptate, 35–120 × 3–6 µm; conidia 50–100 µm long; on Justicia betonica, Asia (India) ……….. C. justiciigena

    Stromata smaller, 10–25 µm diam; conidiophores much shorter, 10–40 × 2.5–5 µm, 0–1-septate; conidia 25–80 µm long; on Justicia spicigera, Central America (Guatemala) ……….. C. jacobiniicola

  • Lepidagathis

    A single species ……….. C. lepidagathidis

  • Pachystachys

    A single species ……….. C. justiciicola

  • Peristrophe

    A single species ……….. C. peristrophes

  • Rhytiglossa

    A single species ……….. C. diantherae

  • Ruellia

    A single species ……….. C. ruellina

  • Rungia

    A single species ……….. C. justiciicola

  • Strobilanthes

    A single species ……….. C. strobilanthis

  • Thunbergia

    1 Conidia acicular to somewhat obclavate, 40–360 × (2−)3–8 µm, base truncate to somewhat obconically truncate ……….. C. thunbergiana

    Conidia acicular, narrower, 45–155 × 2–3 µm, base truncate ……….. C. thunbergiigena

Cercospora species on Acanthaceae

Cercospora acanthi Pass., in Rabenh., Fungi Eur. Exs., Edn Nov., Ser. Sec., Cent. 3 (Resp. Cent. 23), no. 2273 (1876). (Fig. 1)

Fig. 1
figure1

Cercospora acanthi (Rabenh., Fungi Eur. Exs. 2273, HAL, lectotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Synonyms: Cercosporella acanthi (Pass.) D. Sacc., Suppl. Micol. ‘Flora Veneta Critt.’ I: 93 (1899).

Cercosporella compacta Traverso, Hedwigia 43: 422 (1904) [lectotype (designated by Crous & Braun 2003): Italy: Padova, botanical garden, Jun. 1904, A. Pigal [P. Sacc., Mycoth. Ital. 1580] (B 700016201); isolectotypes: Sacc., Mycoth. Ital. 1580, e.g. BPI 420644].

Cercospora acanthi-longifolii Săvul. & Sandu, Acad. Române Mem. Secţ. Şti., ser. 3, 15: 484 (1941) [holotype: Romania: Distr. Severin, near Cazane, on Acanthus longifolius, 16 Jul. 1937 (BUCM)].

Literature: Saccardo (1886: 448; 1906: 562; 1972: 1367), Lindau (1910: 133), Chupp (1954: 21), Braun (1995: 100), Crous & Braun (2003: 40).

Illustration: Chupp (1954: 21, fig. 1).

Exsiccatae: Rabenh., Fungi Eur. Exs. 2273. D. Sacc., Mycoth. Ital. 191, 1580.

Description: Leaf spots amphigenous, circular, subcircular to angular-irregular, 1–12 mm diam, sometimes confluent and larger, sometimes zonate, at first pale greenish, later yellowish to ochraceous, brown, finally greyish brown to greyish white, usually with a narrow darker border, occasionally somewhat raised. Caespituli amphigenous, punctiform, dark brown, later greyish white by abundant conidiation, scattered. Mycelium internal. Stromata well-developed, 10–70 µm diam, substomatal to immersed, large stromata often rupturing the stomata, somewhat erumpent, pale, subhyaline to yellowish brown, later dark brown, composed of swollen hyphal cells, 2.5–10 µm diam, circular to somewhat irregular in shape. Conidiophores in small to large fascicles, dense, arising from stromata, through stomata or erumpent, erect, straight to curved or somewhat geniculate-sinuous, unbranched, 10–40(− 60) × 3–7 µm, 0–2-septate, subhyaline, yellowish to pale olivaceous, paler towards the tip, somewhat darker in mass, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 µm long, mostly with a single or two, occasionally several conspicuous conidiogenous loci, 1.5–2.5 µm diam. Conidia solitary, acicular, smaller conidia narrowly obclavate-cylindrical, straight to curved, 20–130 × 2–5 µm, rarely longer, shorter conidia 1–5-septate, longer ones indistinctly pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate to short obconically truncate, (1.5−)2–2.5(− 3) µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202779): Italy: Parma, Botanical Garden, on Acanthus spinosus, Jul. 1875, G. Passerini [Rabenh., Fungi Eur. Exs. 2273] (HAL). Isolectotypes: e.g., B, BPI 432370, HBG, LE 40368.

Host range and distribution: On Acanthus (hungaricus [balcanicus], mollis [longifolius, niger], spinosus [spinosissimus], Acanthus spp., Ruttya fruticosa), Acanthaceae, Africa (Algeria, Ethiopia), Europe (Germany, Hungary, Italy, Romania).

Notes: The first valid description of Cercospora acanthi dates back to 1876 (in Rabenh., Fungi Eur. Exs. 2273), with a brief description on the label. The secondary description in “Hedwigia 16: 123 (1877)”, usually cited as the original description, is younger; this is a consequence of Saccardo not accepting publication in exsiccate labels as acceptable. This species is a true Cercospora readily distinguishable from the C. apii s. lat. complex by its short, aseptate or sparingly septate, pale conidiophores and acicular to obclavate conidia. Records of this species on Peristrophe bicalyculata from Myanmar (Thaung 1984) are very doubtful and excluded.

Cercospora adhatodae S. Chowdhury, Lloydia 18: 84 “1955” (1956). (Fig. 2)

Fig. 2
figure2

Cercospora adhatodae (K(M) IMI 135861). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Vasudeva (1963: 32), Crous & Braun (2003: 44), Kamal (2010: 13).

Illustration: Chowdhury (1956: 85, fig. 1).

Description: Leaf spots amphigenous, 0.5–8 mm diam, often confluent, forming larger patches, subcircular to irregularly shaped, greyish white. Caespituli amphigenous, fine, dark. Mycelium internal. Stromata lacking or small, substomatal, brown. Conidiophores in small, divergent fascicles, arising from internal hyphae or stromatic hyphal aggregations, through stomata or erumpent, erect, straight, subcylindrical to geniculate in the upper half, 40–190 × 4–6 µm, 0–7-septate, brown to dark brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, about 2–3 µm wide, thickened and darkened. Conidia solitary, acicular, straight to curved, 35–250 × 2.5–5.5 µm, 2–22-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, about 2–3 µm wide, hila thickened and darkened.

Holotype: India: Asom (Assam): Kokilamukh, on Justicia adhatoda, 12 Feb. 1951, S. Chowdhury (HCIO).

Host range and distribution: On Justicia adhatoda [Adhatoda vasica], Acanthaceae, Asia (India, Asom, Karnataka, Uttar Pradesh, West Bengal).

Notes: This is a species of the Cercospora apii s. lat. complex. Several Indian collections from Karnataka, West Bengal and Uttar Pradesh have been examined (K(M) IMI 135861, 293593 and 330436).

Cercospora andrographidicola S.Q. Chen & P.K. Chi, J. S. China Agric. Univ. 11 (3): 61 (1990); as “andrographicola”. (Fig. 3)

Fig. 3
figure3

Cercospora andrographidicola (based on Chi 1994: 94, fig. 86). A. Conidiophore fascicle. B. Conidium. Bar = 10 µm.

Literature: Chi (1994: 94).

Illustration: Chi (1994: 94, fig. 86).

Description: Leaf spots amphigenous, circular, elliptical to irregular, centre pale, margin indistinct, greyish green below. Caespituli amphigenous. Mycelium internal. Stromata well-developed, 16–57 µm diam, brown. Conidiophores in loose fascicles, 4–15, arising from stromata, erect, straight, subcylindrical, non-geniculate, unbranched, 150–233 × 3.3–6.7 µm, 2–10-septate, olivaceous; conidiogenous cells integrated, terminal, usually with a single terminal conidiogenous locus, thickened and darkened. Conidia solitary, acicular, straight to somewhat curved, 200–240 × 3.3–5 µm, hyaline, thin-walled, smooth, apex pointed, base truncate, hila thickened and darkened.

Holotype: China: Guangdong: Gaoyao, on Andrographis paniculata, Oct. 1986, S. G. Chen 123 (Hb. S. China Agric. Univ., Guangzhou).

Host range and distribution: On Andrographis paniculata, Acanthaceae, Asia (China).

Notes: Belonging to the Cercospora apii s. lat. complex, but type material was not available for a re-examination. To-anun et al. (2011) described and illustrated “C. andrographidicola” on Andrographis paniculata from Thailand. This material is, however, morphologically dinstinct by lacking or small stromata, up to 30 µm diam, shorter, geniculate conidiophores, and much shorter, acicular to obclavate conidia, and rather belongs to C. andrographidis.

Cercospora andrographidis Thirum. & Govindu, Sydowia 7: 310 (1953). (Fig. 4)

Fig. 4
figure4

Cercospora andrographidis (BPI 432654). A. Conidiophore fascicle. B. Solitary conidiophore. C. Conidia. Bar = 10 µm.

Literature: Vasudeva (1963: 36), Braun & Crous (2003: 57), Kamal (2010: 16), To-anun et al. (2011: 30), as “C. andrographidicola”.

Illustrations: Thirumalachar & Govindu (1953: plate VI, figs 56), To-anun et al. (2011: 30, fig. 12), as “C. andrographidicola”.

Fig. 5
figure5

Cercospora asystasiana (PC, holotype). A. Conidiophore fascicle. B. Conidiophore, C. Conidia. Bar = 10 µm.

Fig. 6
figure6

Cercospora balaghatensis (K(M) IMI 150920a, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Description: Leaf spots amphigenous, circular to somewhat angular-irregular, 2–6 mm diam, at first dingly greenish, olivaceous or brown, later greyish white, surrounded by a narrow brown margin or marginal line, slightly raised or margin broader, pinkish brown. Caespituli amphigenous, mainly epiphyllous, not very conspicuous. Mycelium internal. Stromata lacking or inconspicuous and small, only a few swollen hyphal cells, 5–20 µm diam, usually intraepidermal, if present to 30 µm diam, brown, cells 3–8 µm diam, wall slightly thickened. Conidiophores solitary or in small divergent fascicles, arising from internal hyphae or aggregations of swollen hyphal cells, erumpent, erect, straight to geniculate, unbranched, 15–165 × 3–6.5 µm, 1–9-septate, light to medium dark brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 15–30 µm long, sympodial, conidiogenous loci conspicuous, thickened and darkened, 2–3 µm diam. Conidia solitary, acicular to somewhat obclavate, straight to curved, 30–150 × 2–4 µm, 1–16-septate, hyaline, thin-walled, smooth, apex subacute, base truncate to somewhat obconically truncate, 2–2.5 µm wide, hila thickened and darkened.

Holotype: India: Bihar: Patna, on Andrographis sp., 20 Jan. 1951, M. J. Thirumalachar (not traced).

Host range and distribution: On Andrographis (paniculata, Andrographis sp.), Acanthaceae, Asia (India, Bihar, Andhra Pradesh, West Bengal; Thailand).

Note: The conidia of this species were described to be “acicular” with obconically truncate base, i.e. they are, at least partly, obclavate as depicted in Thirumalachar & Govindu (1953: plate VI, fig. 6). This could be confirmed on the base of an examined collection from India (West Bengal, Midnapur, Daspur, 20 Mar. 1967, M. Mandal, BPI 432654). To-anun et al. (2011) described and illustrated “C. andrographidicola” on Andrographis paniculata from Thailand. This material, characterised by lacking or small stromata and acicular to obclavate conidia, is quite distinct from true C. andrographidicola and rather belongs to C. andrographidis. An additional sample from India (Daspur, BPI 432654) has been examined.

Cercospora asystasiana J.M. Yen, Rev. Mycol. 32: 180 (1967). (Fig. 5)

Literature: Yen & Lim (1980: 155), Crous & Braun (2003: 70), Nakashima et al. (2010).

Illustration: Yen (1967: 181, fig. 2), Yen & Lim (1980: 206, fig. 9).

Description: Leaf spots amphigenous, on faded leaves, scattered, subcircular, 2–5 mm diam, brown, margin indefinite. Caespituli hypophyllous, rather inconspicuous. Mycelium internal. Stromata lacking or almost so. Conidiophores solitary or in small divergent fascicles, 2–5, arising from internal hyphae or small substomatal hyphal aggregations, through stomata, erect, straight to somewhat curved, subcylindrical to distinctly geniculate-sinuous, unbranched, 30–120(− 135) × 3–6 µm, 0–5-septate, brown to dark brown, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, about 15–30 µm long, conidiogenous loci thickened and darkened, about 2–2.5 µm diam. Conidia solitary, acicular to somewhat obclavate, straight to curved or somewhat sigmoid, 45–185 × 2.5–5 µm, 4–20-septate, hyaline, thin-walled, smooth, apex subacute, base truncate or slightly obconically truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: Singapore: Botanical Garden, on Asystasia chelonoides, 5 Jan. 1966, J. M. Yen 769 (PC).

Host range and distribution: On Asystasia (chelonoides, nemorum), Acanthaceae, known from the type collection [records of Cercospora justiciicola on Asystasia gangetica and C. cf. malloti on A. salicifolia might belong to this species — see notes].

Notes: This species is part of the Cercospora apii s. lat. complex. Records of Cercospora justiciicola on Asystasia gangetica [coromandeliana] (see Cous & Braun 2003: 234) are unclear but might belong to C. asystasiana.

Nguanhom et al. (2015) examined Cercospora species from northern Thailand using molecular methods. Sequences derived from several collections on various unrelated host species, including Asystasia salicifiolia, clustered in a clade tentatively denominated as C. cf. malloti in Groenewald et al. (2013: 157). Taxa belonging to this clade represented the most common Cercospora encounted in this study. Cercospora cf. malloti is morphologically part of the C. apii complex. The true C. malloti was based on North American Cercospora material infecting Mallotus japonicus. The tentative allocation of this clade comprising plurivorous C. apii-like races to C. malloti is neither settled nor finally proven since cultures and sequences based on North American collections retrieved from Mallotus are not yet available. Due to the wide but hitherto little known host range of this taxon, it is not yet possible to exclude that several other host species and older species names of Cercospora might be involved. Therefore, it is currently impossible to resolve the clade concerned.

Cercospora balaghatensis S.M. Singh, Indian Phytopathol. 29: 17 “1976” (1977). (Fig. 6)

Literature: Crous & Braun (2003: 74), Kamal (2010: 20).

Illustration: Singh (1977: 18, fig. 1).

Description: Leaf spots dingy grey with light to dark brown irregular margin. Caespituli amphigenous, scattered, punctiform, dark. Mycelium internal. Stromata 20–85 µm diam, immersed, olivaceous-brown. Conidiophores in large, loose to usually dense fascicles, arising from stromata, erect, straight, subcylindrical, barely geniculate, unbranched, about 10–40 × 3–5 µm, 0–4-septate, pale olivaceous to brownish, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened, about 2–2.5 µm diam. Conidia solitary, acicular or subcylindrical, straight to slightly curved, 20–50 × 2–4 µm, 0–5-septate, hyaline, thin-walled, smooth, apex subacute or subobtuse, base truncate, 1.5–3 µm wide, hila thickened and darkened.

Holotype: India: Madhya Pradesh: Balaghat, on Justicia sp., Acanthaceae, Jun. 1970, S. M. Singh (K(M) IMI 150920a).

Host range and distribution: Only known from the type collection.

Notes: A true Cercospora s. str. distinct from C. apii s. lat. by having very large stromata, short, densely fasciculate conidiophores and short, 0–4-septate conidia.

Cercospora barleriicola Payak & Thirum., Indian Phytopathol. 2: 191 (1949); as “barlericola” (Fig. 7)

Fig. 7
figure7

Cercospora barleriicola (K(M) IMI 102433). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Synonym: Cercospora barleriae-cristatae Govindu & Thirum., Sydowia 10: 273 “1956” (1957) [holotype: India: Karnataka: Bangalore, on Barleria cristata, 10 Dec. 1953, H. C. Govindu (not traced)].

Literature: Chupp (1954: 22), Vasudeva (1963: 46–47), Crous & Braun (2003: 76), Kamal (2010: 21), Meeboon et al. (2007a,b), To-anun et al. (2011: 31)

Illustrations: Govindu & Thirumalachar (1957: plate VIII, fig. 10), Vasudeva (1963: 47, fig. 16), To-anun et al. (2011: 31, fig. 13).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 2–8 mm diam, diffuse discolorations, yellowish to dark reddish brown, later pale brownish to greyish brown or greyish white with darker border. Caespituli amphigenous, subeffuse to punctiform, dark brown. Mycelium internal. Stromata almost lacking or small to moderately large, 10–60 µm diam, olivaceous-brown to dark brown, substomatal to immersed. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to moderately geniculate-sinuous, unbranched, 10–210 × 2.5–6 µm, 1- to pluriseptate throughout, pale to medium olivaceous-brown or brown throughout or paler towards the tip, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, (1.5−)2–3 µm diam. Conidia solitary, longer conidia acicular with truncate base, shorter ones may be obclavate-cylindrical with obconically truncate base, straight to curved, 30–220 × 2–4 µm, 3–18-septate, hyaline, thin-walled, smooth, apex pointed, base truncate to somewhat obconically truncate, 1.5–3 µm wide, hila thickened and darkened.

Holotype: India: Uttar Pradesh: Varanasi, Baranares Hindu University, on Barleria cristata, 9 Dec. 1949, M. M. Payak (not traced).

Host range and distribution: On Barleria (cristata, prionitis, Barleria sp.), Acanthaceae, Asia (India, Jammu and Kashmir, Karnataka, Maharashtra, Madhya Pradesh, Uttar Pradesh; Thailand), West Indies (Jamaica).

Notes: Several Indian collections and a sample from Jamaica on Barleria spp. (K(M) IMI 102433, 163693, 226980, 265815) have been examined and proved to belong to a single variable Cercospora s. str. species belonging to the C. apii s. lat. complex. Since phylogenetic data are not yet available, it remains unclear if a single polymorphous species or several cryptic species are involved.

Cercospora blepharidicola U. Braun, nom. nov. MycoBank MB814564 (Fig. 8)

Fig. 8
figure8

Cercospora blepharidicola (based on Dubay et al. 2011: 516, fig. 3). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora blepharidis R.K. Dubey et al., J. Mycol. Pl. Pathol. 41: 514 (2011), as “blepharia”, nom. illeg. (Art. 53.1), non C. blepharidis Chidd, 1960.

Illustration: Dubey et al. (2011: 516, fig. 3).

Description: Leaf spots amphigenous, mainly epiphyllous, small to large, scattered, dark brown. Caespituli amphigenous, effuse, uniformly distributed. Mycelium internal. Stromata well-developed, to 45 µm diam, dark olivaceous. Conidiophores in loose fascicles, erect, straight, subcylindrical, flexuous, unbranched, arising from a swollen base, about 15–65 × 4–6 µm, 1–3-septate, light olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal, cylindrical, cicatrized, thickened and darkened. Conidia solitary, acicular, straight to curved, 20–112 × 1–4 µm, to 10-septate, hyaline, thin-walled, smooth, apex pointed, base truncate or subtruncate, hila thickened and darkened.

Holotype: India: Madhya Pradesh: North Sagar Forest Division, Toda Khurai, on Blepharis maderaspatensis, Acanthaceae, Dec. 1996, R. K. Dubey AR-58 (HCIO 439724).

Host range and distribution: Only known from the type collection.

Notes: The name C. blepharidis R.K. Dubey et al. is a homonym of C. blepharidis Chidd. (Chiddarwar 1960).

Cercospora crossandrae Jagan., Palan. & Narayans., Madras Agric. J. 59: 672 (1972).

Literature: Crous & Braun (2003: 143), Kamal (2010: 36).

Illustration: Jaganathan et al. (1972: 671, fig. 1).

Description: Leaf spots circular to irregular, 2–5 mm diam, confluent, foliage finally drying up, brown, border yellow, sometimes with concentric rings. Colonies white, later with dark brown margin. Mycelium internal. Conidiophores fasciculate, geniculate, unbranched, about 37.5–62.5 × 6 µm, 4–7-septate, brown. Conidia solitary, acicular to obclavate (“filiform according to the original description), about 53–106 × 3–6 µm, 5–11-septate, hyaline, apex pointed, base truncate to short obconically truncate, probably thickened and darkened.

Holotype: India: Tamil Nadu: Coimbatore, College Orchard of the Tamil Nadu Agricultural University, on Crossandra infundibuliformis, Acanthaceae (T.N. Agric. Univ., Coimbatore, Pl. Pathol. Herb. No. 231).

Host range and distribution: Only known from the type collection.

Notes: It is unknown if type material of this species is maintained. It was not available for examination. Based on the original description and illustration, we suppose that this species belongs to Cercospora s. str. although details of the conidiogenous loci and hila were not described.

Cercospora diantherae Ellis & Kellerm., J. Mycol. 1: 2 (1885). (Fig. 9)

Fig. 9
figure9

Cercospora diantherae (CUP 39680, lectotype). A. Conidiophore fascicles. B. Conidiophores, C. Conidia. Bar = 10 µm.

Synonym: Cercospora jacobinae Mendoza, Philipp. J. Sci. 75: 169 (1941) [holotype: Philippines: Manila, on Justicia carnea, Mendoza, no. 7124 (not traced)].

Literature: Saccardo (1886: 448), Chupp (1954: 23), Crous & Braun (2003: 159).

Illustration: Chupp (1954: 23, fig. 4).

Exsiccatae: Ellis & Everh., Fungi Columb. 695. Ellis & Everh., N. Amer. Fungi 1750. Kellerm. & Swingle, Kansas Fungi 33. Roum., Fungi Sel. Gall. Exs. 5190.

Description: Leaf spots circular to somewhat angular-irregular, 1–5 mm diam, occasionally confluent and larger, slightly zonate, centre greyish white with narrow to often broad brown border. Caespituli amphigenous, dark brown. Mycelium internal. Stromata lacking to small, 10–25 µm diam, composed of a few swollen hyphal cells, brown, substomatal to intraepidermal. Conidiophores in small to moderately large fascicles, 2–15, divergent to moderately dense, arising from internal hyphae or stromatic hyphal aggregations, through stomata or erumpent, erect, straight, subcylindrical to sinuous or somewhat geniculate, unbranched or rarely branched, 10–100 × 3.5–6 µm, aseptate to pluriseptate throughout, pale to medium brown, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal, 10–25 µm long, with a single to several conidiogenous loci, 2.5–3.5 µm diam. Conidia solitary, acicular or subacicular, straight to curved or occasionally somewhat sigmoid, 40–160(− 180) × 2–4 µm, pluriseptate, distance between septa 5–15 µm, hyaline, thin-walled, smooth, apex acute to subobtuse, base usually truncate, occasionally somewhat obconically truncate, 2–3 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202780): USA: Kansas: on Justicia americana, Sep. 1884, W. A. Kellerman (CUP 39680). Isolectotype: PH 1798.

Host range and distribution: On Justicia (americana, carnea, ovata, Justicia sp.), Rhytiglossa humilis [Justicia humilis], Acanthaceae, Asia (Philippines), North America (USA, Delaware, Florida, Illinois, Indiana, Kansas, Maryland, Missouri, Oklahoma, Texas, Washington, West Virginia), South America (Venezuela).

Notes: This species belongs to the Cercospora apii (s. lat.) complex. Type material (collected in Sep. 1884) is not preserved at NY. There are numerous topotypes from 1886, e.g. B; BPI 435404, 435692, 435700, 435701; NY 270701. However, the type material collected in Sep. 1884 was traced in CUP and FH. The CUP material is designated as lectotype. Records of C. diantherae on Jacobinia spp. are doubtful. The hosts concerned probably refer to Justicia spp. in the current sense.

Cercospora hygrophilae Ponnappa, Proc. Indian Acad. Sci., sect. B, 67: 31 (1968). (Fig. 10)

Fig. 10
figure10

Cercospora hygrophilae (K(M) IMI 126157). A. Conidiophores. B. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 222), Kamal (2010: 52).

Illustration: Ponnappa (1968: 32, fig. 1).

Description: Leaf spots amphigenous, often marginal, oblong, oval to irregular, 5–15 mm diam, dark brown to blackish. Caespituli amphigenous, fine, dark. Mycelium internal. Stromata lacking or small, formed as aggregations of some swollen hyphal cells, 10–30 µm diam, substomatal or immersed, globose to somewhat irregular, brown.

Conidiophores in divergent to moderately dense fascicles, 2–15, arising from internal hyphae or stromatic hyphal aggegations, erumpent or through stomata, erect, straight, subcylindrical to somewhat geniculate above, unbranched, 30–500 × 3–8 µm, 1- to pluriseptate throughout, brown, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, about 10–35 µm long, conidiogenous loci conspicuous, thickened and darkened, 2.5–4 µm diam. Conidia solitary, acicular, straight, curved to sigmoid, 50–250 × 2–5(− 6) p m, 5–22-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, 2–4 µm wide, hila thickened and darkened.

Holotype: India: Karnataka: Bannerghatta, on Hygrophila auriculata, 21 Feb. 1967, K. M. Ponnappa (K(M) IMI 126157).

Host range and distribution: On Hygrophila (auriculata [spinosa, Asteracantha longifolia], polysperma, ringens [quadrivalvis]), Acanthaceae, Asia (India, Karnataka, West Bengal).

Notes: This species is a typical member of Cercospora s. str. belonging to the C. apii s. lat. complex, but the conidiophores are very long, 30–500 × 3–8 µm. The original description is misleading and possibly based on not fully developed conidiophores and conidia.

Cercospora jacobiniicola A.S. Mull. & Chupp, Ceiba 1: 174 (1950); as “jacobinicola”. (Fig. 11)

Fig. 11
figure11

Cercospora jacobiniicola (CUP 40082, holotype). A. Conidiophore fascicles. B. Conidiophore, C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 24).

Description: Leaf spots amphigenous, subcircular to irregular, 2–6 mm diam, lead-coloured to blackish, with somewhat raised narrow border line. Caespituli amphigenous, mainly hypophyllous, punctiform, brown. Mycelium internal. Stromata lacking or small, substomatal, about 10–25 µm diam, dark olivaceous to brown. Conidiophores in small to moderately large fascicles, divergent to dense, arising from internal hyphae or stromata, through stomata, erect, straight, subcylindrical-conical to slightly geniculate-sinuous, unbranched, 10–40 × 2.5–5 µm, 0–1-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth; conidiophores reduced to conidiogenous cells or conidiogenous cells integrated, terminal, 10–30 µm long, conidiogenous loci conspicuous, formed as minute circles, about 1 µm diam, only margin slightly thickened and darkened (paracercosporoid). Conidia solitary, cylindrical to obclavate-subcylindrical, straight to slightly curved, 25–80(− 105) × (2.5−)3–4(− 5) µm, 2–7(− 9)-septate, hyaline to very pale greenish or olivaceous, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, occasionally truncate, 1–1.5 µm wide, barely thickened and darkened.

Holotype: Guatemala: Chimaltenango, on Justicia spicigera [Jacobinia spicigera], Acanthaceae, 2 Oct. 1941, A. S. Muller 42 (CUP 40082).

Host range and distribution: Only known from the type collection.

Note: Tentatively maintained in Cercospora. The conidia are colourless or almost so, the conidiogenous loci are conspicuous, but minute, ca. 1 µm wide, and somewhat paracercospora-like. Cultures and results based on molecular sequence analyses are necessary to resolve the true generic affinity of this species.

Cercospora justiciae-adhatodae U. Braun, sp. nov. MycoBank MB814556 (Fig. 12a,b)

Fig. 12a
figure12a

Cercospora justiciae-adhatodae (BPI 1103659, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Fig. 12b
figure12b

Cercospora justiciae-adhatodae (BPI 1103659, holotype). Symptoms on leaves.

Diagnosis: Differs from Cercospora adhatodae and all other Cercospora species on hosts of the Acanthaceae in forming small catenate conidia, (8−)12–35(− 40) × 2–3 µm, (0−)1–3(− 4)-septate.

Description: Leaf spots amphigenous, circular, subcircular to slightly angular-irregular, 0.5–4 mm diam, at first brown, but soon turning greyish white to white, margin narrow, somewhat raised, dark, brown, dark violet to almost blackish. Caespituli amphigenous, punctiform, scattered to dense, brown to dark brown. Mycelium internal. Stromata lacking or almost so to well-developed, 10–30 µm diam, substomatal to intraepidermal, brown, cells 2–5 µm diam, wall at first thin, later slightly thickened. Conidiophores in small, loose to moderately large and dense fascicles, arising from substomatal or intraepidermal hyphae or stromata, emerging through stomata or erumpent, erect, straight to curved, subcylindrical or somewhat attenuated towards the tip to moderately geniculate-sinuous, unbranched, 10–50 × 2–5 µm, 0–3-septate, pale olivaceous to olivaceous-brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, proliferation sympodial, conidiogenous loci conspicuous, 1–2 µm wide, somewhat thickened and darkened. Conidia solitary and catenate, in simple or occasionally branched chains, narrowly cylindrical-fusiform, short obclavate, (8−)12–35(− 40) × 2–3 µm, (0−)1–3(− 4)-septate, hyaline, thin-walled, smooth, apex subacute, subobtuse or short conically truncate, base subtruncate to short obconically truncate, 1–1.5 − m wide, hila slightly thickened and darkened.

Holotype: India: Karnataka: Chikkaballapur District, Nandi Hills, on Justicia adhatoda [Adhatoda vasica], Acanthaceae, 10 Jan. 1969, V. S. Seshadri & K. A. Lucy Channnamma (BPI 1103659).

Host range and distribution: Only known from the type collection.

Notes: Type material of this species was originally deposited as Cercospora adhatodae, but it is quite distinct from the latter species, which belongs to the C. apii complex, by much shorter, narrower, usually 1–3-septate conidia formed in chains. These characteristics distinguish the new species from all other Cercospora species described on hosts belonging to the Acanthaceae. Previously one were inclined to put this species in Passalora s. lat. (including Phaeoramularia), but catenate, colourless conidia are rather in favour of Cercospora s. str., which has recently been shown in the course of phylogenetic studies of cercosporoid hyphomycetes (see Braun et al. 2013). Therefore, the present new species is, at least for the interim, placed in Cercospora although cultures and molecular data are not yet available.

Cercospora justiciigena U. Braun, nom. nov. MycoBank MB814565 (Fig. 13)

Fig. 13
figure13

Cercospora justiciigena (based on Karan & Manoharachary 1978: 158, fig. 1–2). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora acanthacearum var. macrospora Karan & Manohar., Botanique 7: 159 “1976” (1978), non Cercospora macrospora Bres., 1886.

Literature: Crous & Braun (2003: 40), Kamal (2010: 12).

Illustration: Karan & Manoharachary (1978: 158, fig. 1–2).

Description: Leaf spots epiphyllous, zonate, 4–6 mm diam, white, surrounded by a pinkish border. Mycelium internal. Stromata about 30–50 µm diam, dark brown, pseudoparenchymatic. Conidiophores fasciculate, arising from stromata, erect, straight to curved, unbranched, geniculate in the upper half, about 35–120 × 3–6 µm, aseptate, brown, colourless towards the tip; conidiophores reduced to conidiogenous cells, apex rounded with dark annular conidiogenous loci. Conidia solitary, obclavate, straight, about 50–100 × 3–4.5 µm, pluriseptate, hyaline, thin-walled, smooth, tips pointed, base short obconically truncate, hila somewhat thickened and darkened.

Holotype: India: Andhra Pradesh: Hyderabad, University, green house, on Justicia betonica, Acanthaceae, 10 Dec. 1964, D. Karan & C. Manoharachary (HY 197).

Host range and distribution: Only known from the type collection.

Notes: This fungus was introduced as variety of C. acanthacearum, although quite different from the latter species by its much longer conidiophores and much wider conidia. Owing to the description and illustration of conspicuous, dark conidiogenous loci and hila as well as colourless conidia formed singly, C. acanthacearum var. macrospora is undoubtedly a true Cercospora (s. str.) species. The generic affinity of C. acanthacearum is unclear, but this species might rather be a member of Pseudocercospora.

Cercospora justiciicola F.L. Tai, Lloydia 11: 47 (1948); as “justiciaecola”. (Fig. 14)

Fig. 14
figure14

Cercospora justiciicola (based on Guo et al. 2005: 17, fig. 1). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 24), Ellis (1976: 244), Guo et al. (2005: 17), Kamal (2010: 55).

Illustrations: Tai (1948: 44, fig. 9), Ellis (1976: 243, fig. 183 A), Guo et al. (2005: 17, fig. 1).

Description: Leaf spots amphigenous, subcircular, 3–10 mm diam, yellowish, ochraceous, pale brown to finally greyish white, often somewhat zonate, sometimes with yellowish halo. Caespituli amphigenous, punctiform, brown. Mycelium internal. Stromata almost lacking to developed, 10–30 µm diam, brown, substomatal to immersed. Conidiophores in small to moderately large fascicles, divergent, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to usually geniculate-sinuous, unbranched, 25–195 × 3–6 µm, 2–10-septate, olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal to intercalary, about 15–35 µm long, conidiogenous loci thickened and darkened, 1.5–3 µm diam. Conidia solitary, acicular to obclavate-subcylindrical, straight to curved, (15−)40–140(− 155) × 3–4.5(− 5) µm, (2−)3–10(− 17)-septate, hyaline, thin-walled, apex subacute, base truncate to short obconically truncate, 2–3 µm wide, hila thickened and darkened.

Holotype: China: Sichuan: Chengtu, on Justicia sp., 15 Oct. 1944, H. C. Lin (HMAS 12127).

Host range and distribution: On Justicia (betonica, carnea [Jacobinia obtusior], diffusa, flava, gendarussa, procumbens, simplex, Justicia sp.), Pachystachys lutea [Justicia lutea], Rungia (pectinata [Justicia pectinata, Rungia parviflora], repens [Justicia repens], Rungia sp.), Acanthaceae, Africa (Guinea, Mauritius), Asia (Brunei, China; India, Andhra Pradesh, Madhyar Pradesh, Uttar Pradesh; Nepal), North America (USA, Florida).

Notes: A true Cercospora s. str. close to or conspecific with C. apii s. lat. Records of C. justiciicola on Asystasia gangetica [coromandeliana] (Crous & Braun 2003) are unclear, doubtful and might belong to C. asystasiana

Cercospora lepidagathidis Govindu & Thirum., Sydowia 9: 222 (1955).

Literature: Vasudeva (1963:133–134), Crous & Braun (2003: 248), Kamal (2010: 59).

Illustrations: Govindu & Thirumalachar (1955: plate 3, fig. 3), Vasudeva (1963: 134, fig. 92).

Description: Leaf spots amphigenous, circular or subcircular, 4–7 mm diam, centre greyish white, margin pinkish or brown. Caespituli amphigenous. Mycelium internal. Stromata lacking or small, composed of a few swollen hyphal cells, brown. Conidiophores in well-developed, dense fascicles, erect, mostly unbranched, strongly geniculate-sinuous, 33–183 × 2.8–5.7 µm, 1–6-septate, pale olivaceous to brown, tips colourless; conidiogenous cells integrated, terminal or intercalary, with several conspicuous conidiogenous loci. Conidia solitary, obclavate-cylindrical, straight to slightly curved, 23–170 × 2.8–4.2 µm, 1–14-septate, hyaline, thin-walled, smooth, apex pointed, base short obconically truncate, hila thickened.

Holotype: India: Karnataka, Mysore, Bababudans, on Lepidagathis cuspidata, 20 May 1953, H. C. Govindu (not traced).

Host range and distribution: On Lepidagathis (cuspidata, incurva [hyalina]), Acanthaceae, Asia (India, Karnataka, Maharashtra, Uttar Pradesh; Pakistan).

Notes: In the original publication, Govindu & Thirumalachar (1955) mentioned that type material of this species was deposited at BPI, HCIO and IMI. However, type collections could neither be traced at BPI nor IMI. Based on the original description and illustration, this species is undoubtedly a true Cercospora s. str. Records of this species from West Indies (Puerto Rico, and Virgin Islands) on Lepidagathis alopecuroidea are results of misidentifications and belong to Pseudocercospora lepidagathidis.

Cercospora peristrophes Thirum. & Govindu, Sydowia 7: 47 (1953). (Fig. 15)

Fig. 15
figure15

Cercospora peristrophes (BPI 439429, isolectotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora peristrophigena R.K. Chaudhary et al., J. Living World 2: 38 (1995) [holotype: India: UttarPradesh: Nichlaul, Maharajganj, on Peristrophe bicalyculata, Nov. 1993, S. Narayan (HCIO 41989)].

Cercospora peristrophigena S. Narayan et al., in Rao et al., Sugarcane Pathology, Vol. 1: Fungal Diseases: 82 (1999), nom. inval. (Art. 39.1) and nom. illeg. (Art. 53.1) [holotype: India: Uttar Pradesh: Nichlaul, Maharajganj, on Peristrophe bicalyculata, Nov. 1993, S. Narayan (HCIO 41989); isotype: GPU 5070]; as “peristrophegena”.

Literature: Vasudeva (1963: 162), Crous & Braun (2003: 316), Kamal (2010: 73).

Illustrations: Thirumalachar & Govindu (1953: pl. 2, figs 1112), Vasudeva (1963: 163, fig. 115), Chaudhary et al. (1995: 41, fig. 2), Rao et al. (1999: 83, fig. 11).

Description: Leaf spots amphigenous, circular to somewhat angular-irregular, scattered, 0.5–8 mm diam, yellowish brown, brown to dark brown, centre finally dingy grey to greyish white, margin indefinite, marginal slightly raised or surrounded by a darker border. Caespituli amphigenous, scattered, finely punctiform to effuse, greyish white by abundant conidial formation or brownish. Mycelium internal; hyphae branched, septate, pale olivaceous or brownish. Stromata almost lacking or small, 10–25 µm diam, substomatal, olivaceous-brown or brown. Conidiophores solitary or in small fascicles, 2–7, divergent, arising from internal hyphae or stromata, through stomata, erect, straight, somewhat curved to geniculate-sinuous, unbranched, 30–220 × (2.5−)3–7 µm, 1–16-septate, pale olivaceous to olivaceous brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, 10–35 µm long, conidiogenous loci thickened and darkened, 2.5–3.5 µm diam. Conidia solitary, acicular or subacicular to somewhat obclavate-subcylindrical when shorter, 30–130(− 150) × 2.5–4.5 µm, 1–15-septate, hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate or almost so, 2–3 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202781): India: Bihar: Patna, on Peristrophe bicalyculata, 25 Nov. 1952, M. J. Thirumalachar (K(M) IMI 55521). Isolectotypes: BPI 439429, CUP 40770.

Host range and distribution: On Peristrophe bicalyculata, Acanthaceae, Asia (India, Bihar, Maharashtra, Uttar Pradesh; Myanmar), Africa (Somalia).

Notes: This species belongs to the Cercospora apii s. lat. complex. Records from “Nepal” (Crous & Braun 2003, MycoBank and Index Fungorum) refer to the type of C. peristrophigena (Rao et al. 1999), which is, however, incorrect since this species was described from India (Uttar Pradesh).

Cercospora ruellina R.K. Chaudhary et al., J. Living World 2(2): 43 (1995). (Fig. 16)

Fig. 16
figure16

Cercospora ruellina (based on Rao et al. 1999: 87, fig. 13). A. Conidiophore fascicle. B. C. Conidia. Bar = 10 µm.

Synonym: Cercospora ruelliae S. Narayan et al., in Rao et al., Sugarcane Pathology, Vol. 1: Fungal Diseases: 86 (1999), nom. inval. (Art. 39.1) [holotype: India: Uttar Pradesh: Gorakhpur, Maharajganj, on Ruellia prostrata, Dec. 1993, S. Narayan (HCIO 41994); isotype: GPU 5077].

Literature: Crous & Braun (2003: 360), Kamal (2010: 83).

Illustration: Rao et al. (1999: 87, fig. 13).

Description: Leaf spots amphigenous, circular or almost so, 1–6 mm diam, brown to blackish on the upper leaf surface, whitish green with olivaceous margin below. Caespituli amphigenous, effuse. Mycelium internal; hyphae branched, septate, hyaline to pale olivaceous. Stromata substomatal, 10–35 µm diam, olivaceous to olivaceous-brown. Conidiophores solitary or in small fascicles, 2–5, arising from stromata, through stomata, erect, straight to geniculate-sinuous, unbranched, about 20–125 × 2.5–5 µm, 2–9-septate, light olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary; conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, narrowly acicular, shorter conidia sometimes subcylindrical to somewhat obclavate, straight to curved, often with short lateral germ tubes which may give rise to secondary conidia, about 30–145 × 1.5–2.5 µm, 3–18-septate, hyaline, thin-walled, smooth, apex obtuse to subacute, base truncate, hila thickened and darkened.

Holotype: India: Uttar Pradesh: Gorakhpur, Maharajganj, on Ruellia prostrata, Dec. 1993, S. Narayan (HCIO 41994). Isotype: GPU 5077.

Host range and distribution: Only known from the type collection.

Notes: This species is a typical member of the Cercospora apii s. lat. complex with acicular conidia. A morphologically well agreeing North American sample on Ruellia ciliosa has been examined (USA, Illinois, Chandlerville, 18 Aug. 1886, A. B. Seymour, BPI 435186), although the conidiophores in the latter collection are much longer and somewhat broader, 60–280 × 4–7 µm. However, if Indian and North America collections are conspecific requires phylogenetic examination and confirmation. Indian as well as North American samples seems to pertain to plurivorous species of the C. apii complex.

Cercospora strobilanthis Chidd., Mycopathol. Mycol. Appl. 17: 77 (1962); as “strobilanthidis”. (Fig. 17)

Fig. 17
figure17

Cercospora strobilanthis (K(M) IMI 83190, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Crous & Braun (2003: 390), Guo (2001: 343–344), Guo et al. (2005: 18), Kamal (2010: 89).

Illustration: Chiddarwar (1962: 78, pl. II, figs 13), Guo et al. (2005: 19, fig. 2).

Description: Leafspots amphigenous, circular to irregular, 2–10 mm diam, at first pale greenish, later brownish, finally greyish white or white, with darker border, narrow to moderately wide, brown to almost black. Caespituli hypophyllous, scattered, punctiform, fine, dark. Mycelium internal. Stromata almost absent or small, mainly substomatal, 10–25 µm diam, brown. Conidiophores in divergent fascicles, 2–15, rarely solitary, arising from stromata, through stomata, erect, straight, subcylindrical to distinctly geniculate, 1–6 times, unbranched, 35–250 × 4–5.5 µm, 1–7-septate throughout, pale brown or olivaceous-brown, wall thin or slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened, 1.5–2.5 µm diam. Conidia solitary, acicular to slightly obclavate, straight to curved, 30–195 × 3–5(− 5.5) µm, 3–16-septate, hyaline, thin-walled, smooth, apex pointed, base truncate to usually somewhat attenuated at the very base (very short obconically truncate), 1.5–2.5 µm, hila thickened and darkened.

Holotype: India: Maharashtra: Mumbai, Matheran, on Strobilanthes sp., 12 Dec. 1956, P. P. Chiddarwar 29 (K(M) IMI 83190).

Host range and distribution: On Strobilanthes sp., Acanthaceae, Asia (China, Guangdong; India, Maharashtra).

Note: A true Cercospora s. str. belonging to the C. apii s. lat. complex, but well characterised by conidia that are often somewhat attenuated at the very base.

Cercospora thunbergiana J.M. Yen, Rev. Mycol. 30: 198 (1965); as “thunbergiaena”. (Fig. 18)

Fig. 18
figure18

Cercospora thunbergiana (PC, lectotype). A. Conidiophore fascicle. B. Conidophore tips. C. Conidia. Bar = 10 µm.

Literature: Yen & Lim (1980: 165), Braun & Castañeda Ruiz (1991: 291), Braun et al. (1992: 363), Crous & Braun (2003: 403), Braun & Urtiaga (2008: 485), Kamal (2010: 93), Guo et al. (2005: 18).

Illustrations: Yen (1965: 199, fig. 13), Yen & Lim (1980: 223, fig. 26), Guo et al. (2005: 20, fig. 3).

Description: Leaf spots amphigenous, circular, subcircular or somewhat irregular, 1–7 mm diam, brown, greyish brown, finally greyish white, with dark brown border, sometimes with diffuse brownish halo, finally sometimes with shot-hole symptoms. Caespituli amphigenous, mostly hypophyllous, not very distinct or finely punctiform, dark. Mycelium internal. Stromata lacking or small, forming small aggregations of swollen hyphal cells, 10–30 µm diam, brown. Conidiophores in small fascicles, 2–15, divergent, occasionally solitary, arising from internal hyphae or small stromata, through stomata, erect, straight to geniculate-sinuous, unbranched, 40–220 × (3−)4–8 µm, 2–9-septate, brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, 10–40 µm long, conidiogenous loci conspicuous, thickened and darkened, (1.5−)2–5 µm wide. Conidia solitary, acicular to somewhat obclavate, straight to curved, 40–360 × (2−)3–8 µm, 3–34-septate, occasionally somewhat constricted at septa, hyaline, thin-walled, smooth, base truncate or slightly obconically truncate, 2–5 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202782): Singapore: Bukit Timah (Hwa Chung College), on Thunbergia alata, 3 Aug. 1964, S. H. Yen 73 (PC). Isolectotype: K(M) IMI 120995.

Host range and distribution: On Thunbergia (alata, erecta, grandiflora, Thunbergia sp.), Acanthaceae, Asia (Brunei; India, Andhra Pradesh, Uttar Pradesh; Singapore), South America (Venezuela), West Indies (Cuba).

Cercospora thunbergiigena U. Braun & Crous, nom. nov.

MycoBank MB814567 (Fig. 19)

Fig. 19
figure19

Cercospora thunbergiigena (K(M) IMI 345300, holotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora thunbergiana R.K. Srivast et al., Kavaka 20/21: 42 “1992/1993” (1995), nom. illeg. (Art. 53.1), non C. thunbergiana J.M. Yen, 1965; as “thumbergiana”.

Literature: Crous & Braun (2003: 403), Kamal (2010: 93).

Illustration: Srivastava et al. (1995: 41, fig. 4).

Description: Leaf spots amphigenous, often vein-limited, mostly marginal, 3–10 mm diam, dark brown to blackish. Caespituli amphigenous, fine. Mycelium internal. Stromata absent or small, substomatal, 10–20 µm diam, brown. Conidiophores in small, loose fascicles, arising from small stromata, through stomata, erect, straight to moderately geniculate, unbranched, about 50–220 × 4–5 µm, 3–10-septate, brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened, about 2 µm diam. Conidia solitary, narrowly acicular, straight to curved, 45–155 × 2–3 µm, 4–14-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, 1.5–2 µm wide, hila thickened and darkened.

Holotype: India: Uttar Pradesh: Gorakhpur, Ramgarh area, on Thunbergia grandiflora, Acanthaceae, Dec. 1989, V. P. Pandey (K(M) IMI 345300).

Host range and distribution: Only known from the type collection).

Notes: This species belongs to the C. apii s. lat. complex. It resembles C. thunbergiana, but differs in having much narrower conidia.

Doubtful, excluded and insufficiently known species

Cercospora acanthacearum Govindu & Thirum., Sydowia 8: 221 (1954).

Literature: Vasudeva (1963: 30), Crous & Braun (2003: 40). Kamal (2010: 12).

Illustration: Govindu & Thirumalachar (1954: pl. 6, fig. 1).

Description: Leaf spots circular to irregular, 2–4 mm diam, centre greyish white, surrounded by a pinkish or light brown border. Caespituli mostly epiphyllous. Mycelium internal. Stromata small, only composed of a few brown swollen hyphal cells, 15–30 µm diam. Conidiophores fasciculate, arising from stromatic hyphal aggregations, erect, straight to sinuous, geniculate, unbranched, 14–28.5 × 2.8–4.2 µm, aseptate, pale brown, thin-walled, smooth; conidiophores reduced to conidiogenous cells. Conidia solitary, narrowly obclavate, 14–35.5 × 2–3 µm, 1–6-septate, hyaline, thin-walled, smooth, apex pointed, base short obconically truncate.

Holotype: India: Karnataka: Mysore, Nandi Hills, on Justicia betonica, Acanthaceae, 31 Jan. 1953, H. C. Govindu (not traced).

Host range and distribution: Only known from the type collection.

Notes: According to Govindu & Thirumalachar (1954), type material of new species described in their paper had been deposited at BPI, IMI, and HCIO. However, type material could not be traced, neither in BPI nor in IMI (now K). Details of the structure of the conidiogenous loci were not provided by Govindu & Thirumalachar (1954), but based on the original description this species does probably not belong to Cercospora s. str. It might be a species of Pseudocercospora, but a re-examination of type material or new collections agreeing with the original description are necessary to answer this question.

Cercospora peristrophes E. Castell., Elenco Annotato dei Funghi della Somalia (Biblioteca Agraria Tropicale): 65 (1988), as “peristrophis”, nom. illeg. (Art. 53.1), non C. peristrophes Thirum. & Govindu, 1953.

Literature: Crous & Braun (2003: 316).

Note: Cercospora peristrophes E. Castell. is a homonym of C. peristrophes Thirum. & Govindu, 1953.

Passalora

Key to Passalora species on Acanthaceae
  1. 1

    Conidia solitary, obclavate, short, 25–50 × 3.5–5 µm, 1–3(− 4)-septate; on Barleria lupulina ………. P. barleriigena

    Conidia in chains, either much longer, 20–95 µm, or much narrower, 2–3 µm; on other hosts ……… 2

  2. 2(1)

    Conidiophores 100–325 µm long, pluriseptate; conidia cylindrical or somewhat cylindrical-obclavate, 20–95 × 4–6 µm, 1–6-septate, pale olivaceous; on Acanthus arboreus ……… P. acanthicola

    Conidiophores much shorter, 10–50 µm; conidia 8–40 × 2–3 µm, (0−)1–3(− 4)-septate, hyaline; on Adhatoda vasica ……… see Cercospora justiciae-adhatodae

Passalora species on Acanthaceae

Passalora acanthicola (Hansf.) U. Braun & Crous, Mycosphaerella and Anam. 1: 40 (2003). (Fig. 20)

Fig. 20
figure20

Passalora acanthicola (K(M) IMI 4557a, holotype). A. Base of conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora acanthicola Hansf., Proc. Linn. Soc. London 156: 121 (1944).

Synonym: Phaeoramularia acanthicola (Hansf.) Deighton, Trans. Brit. Mycol. Soc. 88: 385 (1987).

Literature: Chupp (1954: 22).

Illustration: Deighton (1987: 387, fig. 17).

Description: Leaf spots at first diffuse, yellowish, later angular, 5–8 mm diam, brown. Caespituli hypophyllous, effuse, deep reddish brown, more or less vein-limited. Mycelium internal; hyphae 2.5–4 µm wide, colourless. Stromata lacking or almost so. Conidiophores in small loose fascicles, to 10, arising from internal hyphae or small aggregations of swollen hyphal cells, emerging through stomata, erect, geniculate-sinuous, simple or branched, 100–325 × 4–6 µm, pluriseptate throughout, moderately brown or reddish brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened, about 1.5–2 µm diam. Conidia catenate, in simple or branched chains, cylindrical or somewhat obclavate-cylindrical, straight or slightly curved, 20–95 × 4–6 µm, 1–6-septate, pale olivaceous, thin-walled, smooth, apex obtuse or subtruncate, base short obconically truncate, 1.5–2 µm wide, hila thickened and darkened.

Holotype: Uganda: Kampala, Kawandra, on Acanthus arboreus, Acanthaceae, Jan. 1943, C. G. Hansford 3144 (K(M) IMI 4557a).

Host range and distribution: Only known from the type collection.

Passalora barleriigena Meeboon & Hidayat, Mycotaxon 102: 140 (2007). (Fig. 21)

Fig. 21
figure21

Passalora barleriigena (based on Meeboon et al. 2007a: 141, fig. 1). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Meeboon et al. (2007a: 141, fig. 1).

Description: Leaf spots amphigenous, subcircular to irregular, 1–14 mm diam, pale to pale brown, margin at first indefinite, later conspicuous, dark. Caespituli amphigenous. Mycelium internal. Stromata substomatal, small, 7.5–20 µm diam, composed of 3–7 swollen hyphal cells, brown. Conidiophores in dense fascicles, 4–7, arising from stromata, through stomata, erect, straight, subcylindrical, somewhat attenuated towards the tip, unbranched, 1–2 times geniculate, about 20–65 × 3.5–5 µm, 1–3-septate, darker brown below, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci conspicuous, thickened and darkened, 1–2 µm diam. Conidia solitary, mostly obclavate, occasionally oblong cylindrical, straight to occasionaly somewhat curved, 25–50 × 3.5–5 µm, 1–3(− 4)-septate, subhyaline to pale brown, thin-walled, smooth, apex obtuse, base short obconically truncate, 1–2 µm wide, hila thickened and darkened.

Holotype: Thailand: Chiang Mai Province: Doi Suthep-Pui National Park, on Barleria lupulina, 30 Dec. 2005, J. Meeboon (CMU 28212). Paratype: the same locality, 10 Dec. 2006, J. Meeboon & I. Araki (CMU 28213).

Host range and distribution: On Barleria lupulina, Acanthaceae, Asia (Thailand).

Pseudocercospora

Key to Pseudocercospora species on Acanthaceae
  1. 1

    Mycelium internal and external; superficial hyphae with solitary conidiophore in vivo developed ……….. 2

    Mycelium internal; superficial hyphae with solitary conidiophores in vivo not developed ……….. 12

  2. 2(1)

    Stromata lacking; conidiophores 50–130 × 4–6.5 µm; conidia 3.5–6.5 µm long; on Justicia spp ……….. P. justiciae

    Stromata developed; and/or conidiophores shorter, to about 80 µm, and/or narrower, 2–4 µm; and/or conidia much narrower, 1.5–4 µm ……….. 3

  3. 3(2)

    Stromata lacking or almost so ……….. 4

    Stromata developed, 10–40 µm diam ……….. 7

  4. 4(3)

    Conidiophores relatively short, 5–50(− 60) µm ……….. 5

    Conidiophores longer and broader, to 120 × 2.5–6 µm, longer conidiophores with more than two septa (pluriseptate); on other hosts ……….. 6

  5. 5(4)

    Conidiophores short, 5–30 × 1.5–3 µm, 0–1-septate; on Thunbergia spp ……….. P. thunbergiae

    Conidiophores 5–50(− 60) × 2–5 µm, 0–3-septate; on Rhinacanthus nasutus ……….. P. rhinacanthi

  6. 6(4)

    Conidia cylindrical or subcylindrical, base truncate to somewhat obconically truncate; conidiophores to 120 µm long; on Strobilanthes cusia ……….. P. baphiacanthi

    Conidia obclavate-cylindrical, base consistently obconically truncate; conidiophores shorter, to 80 µm long; on Lepidagathis alopecuroidea ……….. P. lepidagathidis

  7. 7(3)

    Conidiophores 10–80 × 3–5 µm, 0–5-septate; conidia rather long, 60–175 × 3.5–5 µm, 4–11-septate, hila 2–2.5 µm wide; on Thunbergia alata ……….. P. thunbergiicola

    Conidiophores shorter, 5–50 × 1.5–5 µm; conidial hila 0.8–2 µm wide; on other hosts ……….. 8

  8. 8(7)

    Conidiophores 20–50 µm long, 2–6-septate; on Justicia japonica ……….. P. justiciicola

    Conidiophores 0–3-septate; on other hosts ……….. 9

  9. 9(8)

    Conidia obclavate-cylindrical to subacicular; conidia with pointed apex; on Acanthus guineensis ……….. P. acanthi

    Conidia obclavate-cylindrical, subacicular conidia lacking; on other hosts ……….. 10

  10. 10(9)

    Conidial base truncate to obconically truncate; on Odontonema callistachyum ……….. P. odontonematis

    Conidial base consistently obconically truncate ……….. 11

  11. 11(10)

    Conidiophores 5–30 × 1.5–4 µm, 0–1(− 2)-septate; on Justicia galapagana ……….. P. consociata var. dimorpha

    Conidiophores to 60 µm long, 0–3-septate; on Rhinacanthus nasutus P. rhinacanthi

  12. 12(1)

    Stromata lacking; conidia 15–65 × 2–4 µm, hila 1–1.5 µm wide; on Blechum pyramidatum ……….. P. blechi

    Stromata developed; and/or conidia longer, to 150 µm, hila broader, 1.5–2.5 µm; on other hosts ……….. 13

  13. 13(12)

    Stromata lacking or almost so; conidiophores 20–130 × 4–7 µm; on Barleria cristata ……….. P. barleriae

    Stromata developed; and/or conidiophores shorter, to 75 µm, and above all narrower, 2–5 µm wide; on other hosts ……….. 14

  14. 14(13)

    Conidiophores short, 5–40 µm, 0–1(− 2)-septate; on Cynarospermum, Dyschoriste, Justicia, Rhinacanthus, Ruellia ……….. 15

    Conidiophores longer, to 125 µm, and/or (if shorter) 0–5-septate 17

  15. 15(14)

    Conidia acicular to obclavate-cylindrical, base truncate to long obconically truncate, 2–2.5 µm wide; on Cynarospermum ……….. P. blepharidis

    Conidia obclavate-cylindrical, acicular conidia not formed, base 1.5–2 µ wide; on other hosts ……….. 16

  16. 16(15)

    Conidiophores 5–30 × 1.5–4 µm, 0–1(− 2)-septate (external mycelium lacking); on Dyschoriste, Justicia, Ruellia ……….. P. consociata var. consociata

    Conidiophores to 60 µm long, 0–3-septate (external mycelium usually developed); on Rhinacanthus nasutus ……….. P. rhinacanthi

  17. 17(14)

    Conidiophores long, 20–125 µm; on Asystasia gangetica ……….. P. asystasiae

    Conidiophores shorter, to 75 µm, average < 50 µm; on other hosts (see “Tabular key to Pseudocercospora species on Acanthaceae according to host genera” — further identification just based on morphology barely possible)

Tabular key to Pseudocercospora species on Acanthaceae according to host genera
  • Acanthus

    A single species ………. P. acanthi

  • Asystasia

    A single species ………. C. asystasiae

  • Baphicacanthus, see Strobilanthes

  • Barleria

    A single species ………. P. barleriae

  • Blechum

    A single species ………. P. blechi

  • Blephris, see Cynarospermum

  • Cynarospermum

    A single species ………. P. blepharidis

  • Dicliptera

    A single species ………. P. diclipterae

  • Dyschoriste

    A single species ………. P. consociata var. consociata

  • Ecbolium

    A single species ………. P. ecbolii

Justicia
  1. 1

    Mycelium internal, superficial hyphae with solitary conidiophores lacking ………. P. consociata var. consociata

    Mycelium internal and external, superficial hyphae with solitary conidiophores developed ………. 2

  2. 2(1)

    Stroma lacking; conidiophores 50–130 × 4–6.5 µm; conidia cylindrical to obclavate-subcylindrical, 40–110 × 3.5–6.5 µm, hila 2–2.5 µm wide ………. P. justiciae

    Stromata developed, 10–25 µm diam; conidiophores shorter and narrower, 20–50 × 1–5 µm; conidia narrower, obclavate, 35–120 × 1.5–5 µm, hila 1–2 µm wide ………. P. justiciicola

  • Lepidagathis

    A single species ………. P. lepidagathidis

  • Odontonema

    A single species ………. P. odontonematis

  • Rhinacanthus

    A single species ………. P. rhinacanthi

  • Ruellia

    A single species ………. P. consociata var. consociata (and ? P. justiciicola)

  • Rungia

    A single species ………. P. rungiae

  • Strobilanthes

    A single species ………. C. baphicacanthi

  • Thunbergia

    1 Stromata lacking; conidiophores short and narrow, 5–30 × 1.5–3 µm; conidia 20–95 × 1.5–4 µm, 0–8-septate, subhyaline, pale yellowish green to very pale olivaceous ………. P. thunbergiae

    Stromata to 30 µm diam; conidiophores longer and broader, 10–80 × 3–5 µm; conidia also longer and broader, 60–175 × 3.5–5 µm, 4–11-septate, pale olivaceous-brown ………. P. thunbergiicola

Pseudocercospora species on Acanthaceae

Pseudocercospora acanthi Deighton, Trans. Brit. Mycol. Soc. 88: 381 (1987). (Fig. 22)

Fig. 22
figure22

Pseudocercospora acanthi (K(M) IMI 7695, holotype). A. Conidiophore fascicles. B. Solitary conidiophore arising from superficial hypha. C. Conidia. Bar = 10 µm.

Illustration: Deighton (1987: 383, fig. 14).

Description: Leaf spots amphigenous, subcircular, to 5 mm diam, grey-brown, with narrow dark brown border, somewhat raised. Caespituli amphigenous rather pale olivaceous, punctiform, scattered. Mycelium internal and external; internal hyphae 2.5–6.5 µm wide, subhyaline, superficial hyphae 1.5–3 µm wide, pale olivaceous, septate, thin-walled, smooth. Stromata well-developed, 20–40 µm diam, compact, brown. Conidiophores in small to sometimes large and dense fascicles, 8–50, arising from stromata, or occasionally solitary, arising from superficial hyphae, erect, straight, subcylindrical-conical to somewhat geniculate-sinuous, unbranched, 20–40 × 3–4.5 µm, 0–1(− 2)-septate, pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 20–30 µm long, conidiogenous loci inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, obclavate-subcylindrical to subacicular, straight to curved or somewhat sigmoid, 30–90 × 2.5–4 µm, 3–8-septate, pale olivaceous, apex pointed, base obconically truncate to almost rounded, 1.5–2 µm wide, hila unthickened, not darkened.

Holotype: Sierra Leone: Gorahun (Tunkia), on Acanthus guineensis, Acanthaceae, 3 Apr. 1939, F. C. Deighton, M 1937 (K(M) IMI 7695).

Host range and distribution: Only known from the type collection.

Pseudocercospora asystasiae (J.M. Yen) J.M. Yen, Gard. Bull., Singapore 33: 169 (1980). (Fig. 23)

Fig. 23
figure23

Pseudocercospora asystasiae (PC, holotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora asystasiae J.M. Yen, Rev. Mycol. 32: 178 (1967).

Literature: Yen & Lim (1980: 169), Crous & Braun (2003: 70), Nakashima et al. (2010).

Illustrations: Yen (1967: 179, fig. 1), Yen & Lim (1980: 228, fig. 31).

Description: Leaf spots amphigenous, subcircular to somewhat irregular, 2–10 mm diam, at first pale greenish, greyish green, later yellowish brown to brownish, greyish brown, finally greyish white, occasionally zonate, margin indistinct to distinct, dark brown to blackish. Caespituli amphigenous, indistinct to punctiform, brown, scattered to confluent and denser. Mycelium internal. Stromata lacking or small, substomatal, 10–30 µm diam, subglobose, brown. Conidiophores in small to moderately large fascicles, loose to moderately dense, arising from stromata, emerging through stomata, erect, straight, subcylindrical to flexuous, sinuous or somewhat geniculate-sinuous, unbranched, 20–125 × 3–5 µm, 0–5-septate, olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal, about 10–40 µm long, conidiogenous loci inconspicuous to subdenticulate, neither thickened nor darkened. Conidia solitary, obclavate-cylindrical, straight to curved, 25–90(− 110) × (2.5−)3–4.5(− 5) µm, 2–8(− 9)-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: Singapore: Botanical Garden, on Asystasia gangetica, 5 Jan. 1966, J. M. Yen 731 (PC).

Host range and distribution: On Asystasia (gangetica [coromandeliana], nemorum), Acanthaceae, Africa (Ivory Coast), Asia (Indonesia, Japan, Singapore).

Note: A second collection on Asystasia gangetica from Ivory Coast (29 Dec. 1974, G. Gilles 47), deposited at PC, has been examined.

Pseudocercospora baphicacanthi W.H. Hsieh & Goh, Bot. Bull. Acad. Sin. 30: 123 (1989). (Fig. 24)

Fig. 24
figure24

Pseudocercospora baphicacanthi (based on Hsieh & Goh 1990: 13, fig. 2). A. Solitary conidiophore, arising from superficial hyphae or emerging through stomata. B. Conidiophores, C. Conidia. Bar = 10 µm.

Literature: Hsieh & Goh (1990: 12), Guo & Hsieh (1995: 1), Guo et al. (1998: 11).

Illustration: Hsieh & Goh (1990: 13, fig. 2), Guo & Hsieh (1995: 2, fig. 1), Guo et al. (1998: 10, fig. 1).

Description: Leaf spots irregularly shaped, 2–12 mm diam, sometimes confluent, forming yellowish patches on the upper leaf surface, yellowish brown below, margin indefinite. Caespituli hypophyllous. Mycelium internal and external; superficial hyphae emerging through stomata. Stromata lacking. Conidiophores solitary or in small, loose fascicles, emerging through stomata or solitary, arising from superficial hyhae, lateral, erect to decumbent, straight to curved, geniculate-sinuous, simple or branched, 20–120 × 3–6 µm, septate, pale brown throughout or paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous or visible as truncate tips, about 2 µm wide, unthickened, not darkened. Conidia solitary, cylindrical or subcylindrical, straight to somewhat curved, 30–100 × 3–4 µm, 2–7-septate, greenish to pale olivaceous, thin-walled, smooth, apex obtuse or subobtuse, base truncate to short obconically truncate, hila unthickened, not darkened.

Holotype: Taiwan: Hsitou, Nantou Hsien, on Strobilanthes cusia [Baphicacanthus cusia], Acanthaceae, 28 Mar. 1985, W. H. Hsieh (NCHUPP-162). Isotype: K(M) IMI 312069.

Host range and distribution: Only known from the type collection.

Pseudocercospora barleriae (J.M. Yen & Lim) U. Braun, Fungal Diversity 8: 60 (2001). (Fig. 25)

Fig. 25
figure25

Pseudocercospora barleriae (PC, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora barleriae J.M. Yen & Lim, Cah. Pacifique 17: 100 (1973).

Literature: Yen & Lim (1980: 156), Crous & Braun (2003: 76).

Illustrations: Yen & Lim (1973: 110, fig. 3; 1980: 207, fig. 10), Braun (2001: 58, fig. 15).

Description: Leaf spots amphigenous, angular, vein-limited, 1–5 mm diam or confluent and larger, blackish on the upper leaf surface, dark brown below, sometimes forming large blackish brown patches, covering large leaf segments. Caespituli hypophyllous, effuse, dark brown, sometimes velvety. Mycelium internal. Stromata lacking or almost so. Conidiophores in small to moderately large fascicles, loose, arising from internal hyphal or small hyphal aggregations, through stomata, erect to decumbent, flexuous, simple or occasionally branched, somewhat geniculate-sinuous, 20–130 × 4–7 µm, (0−)1–7-septate, olivaceous to olivaceous-brown, tips paler, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–40 µm long, conidiogenous loci inconspicuous or subconspicuous by being subdenticulate or somewhat refractive, but neither thickened nor darkened. Conidia solitary, obclavate to obclavate-cylindrical, straight to somewhat curved, (10−)15–80 × 4–5 µm, 1–8-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex subacute to obtuse, base obconically truncate, 2–2.5 µm wide, hila unthickened, not darkened.

Holotype: Singapore: on Barleria cristata, 3 Mar. 1972, G. Lim 73 (PC). Topotype: June 1972 (PC).

Host range and distribution: On Barleria cristata, Acanthaceae, Asia (Singapore).

Notes: Yen & Lim (1980) retained this species in Cercospora and described “brown scars”. However, the examination of type material showed that this species has to be reallocated to Pseudocercospora.

Pseudocercospora blechi U. Braun et al., Feddes Repert. 113: 120 (2002). (Fig. 26)

Fig. 26
figure26

Pseudocercospora blechi (CUP-VZ-002240, holotype). A. Conidiophore fascicles. B. Conidiophore. C. Conidia. Bar = 10 µm.

Synonym: Cercospora blechi Chupp & A.S. Mull., Bol. Soc. Venez. Ci. Nat. 8: 37 (1942), nom. inval. (Art. 39.1).

Literature: Chupp (1954: 22), Crous & Braun (2003: 82).

Illustrations: Chupp (1954: 21, fig. 2), Braun et al. (2002: 121, fig. 8).

Description: Leaf spots mainly epiphyllous, indistinct to irregular and somewhat reddish, 0.5–3 mm diam, sometimes confluent, margin indefinite. Caespituli hypophyllous, subeffuse, dark olivaceous to blackish. Mycelium internal. Stromata lacking or small, composed of a few swollen hyphal cells, brown. Conidiophores in small, loose fascicles, arising from internal hyhae or hyphal aggregations, emerging through stomata, erect, flexuous, geniculate-sinuous, simple or rarely branched, 10–90 × 2–4 µm, 0–3-septate, pale olivaceous or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, straight to slightly curved, 15–65 × 2–4 µm, 1–6-septate, subhyaline to pale olivaceous, apex obtuse to subacute, base obconically truncate, 1–1.5 µm wide, hila unthickened, not darkened.

Holotype: Venezuela: Caracas, on Blechum pyramidatum, 28 Jul. 1938, A. S. Muller 2240 (CUP-VZ-002240). Isotype: VIA.

Host range and distribution: On Blechum pyramidatum [brownei], Acanthaceae, Central and South America (Panama, Venezuela), West Indies (Cuba, Puerto Rico, Virgin Islands).

Pseudocercospora blepharidis (Chidd.) U. Braun & Crous, Mycosphaerella and Anam.: 82 (2003). (Fig. 27)

Fig. 27
figure27

Pseudocercospora blepharidis (K(M) IMI 83163, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora blepharidis Chidd., Sydowia 13: 154 “1959” (1960).

Literature: Kamal (2010: 155).

Illustration: Chiddarwar (1960: plate V, figs 45).

Description: Leaf spots amphigenous, subcircular, ovoid to irregular, 3–18 mm diam, pale to dark brown, sometimes somewhat zonate, above all on the lower leaf surface. Caespituli amphigenous, punctiform, brown. Mycelium internal. Stromata well-developed, substomatal, 20–30 µm diam, olivaceous-brown. Conidiophores in small to moderately large fascicles, arising from stromata, through stomata, divergent to dense, straight, subcylindrical-conical to slightly geniculate-sinuous, unbranched, 5–40 × 3–5 µm, 0–1(− 2)-septate, pale to medium brown or olivaceous-brown, wall thin or only slightly thickened, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–25 µm long, conidiogenous loci inconspicuous. Conidia solitary, acicular to obclavate-cylindrical, straight to curved, 30–80 × 2.5–3.5 µm, 3–8-septate, subyhaline to pale olivaceous, thin-walled, smooth, apex subacute to obtuse, base truncate to long obconically truncate, 2–2.5 µm wide, hila unthickened, not darkened.

Holotype: India: Maharashtra: Khandala, on Cynarospermum asperrimum [Blepharis asperrima], Acanthaceae, 9 Nov. 1956, P. P. Chiddarwar 2 (K(M) IMI 83163).

Host range and distribution: Only known from the type collection.

Pseudocercospora consociata (G. Winter) Y.L. Guo & X.J. Liu, Mycosystema 2: 232 (1989). var. consociata (Fig. 28a)

Fig. 28
figure28

a,b a. Pseudocercospora consociata var. consociata (BPI 435187, neotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. b. P. consociata var. dimorpha (HAL 2190 F, holotype). A. Superficial hyphae. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophore fascicles. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Cercospora consociata G. Winter, Hedwigia 22: 70 (1883).

Literature: Saccardo (1886: 470), Chupp (1954: 23), Katsuki (1965: 7), Guo & Hsieh (1995: 1), Guo et al. (1998: 11), Crous & Braun (2003: 136), Braun & Freire (2006: 236), Kamal (2010: 166), Phengsintham et al. 2013: 110).

Illustration: Chupp (1954: 21, fig. 3), Guo & Hsieh (1995: 3, fig. 2), Guo et al. (1998: 12, fig. 2), Phengsintham et al. 2013: 111, fig. 67).

Exsiccatae: Ellis & Everh., N. Amer. Fungi 2477.

Description: Leaf spots lacking or almost so, inconspicuous to diffuse yellowish to brownish discolorations, or forming subcircular to angular-irregular lesions, 1–10(− 15) mm diam, brownish, finally greyish brown to greyish white, margin indefinite or darker brown, occasionally reddish brown, sometimes slightly raised, narrow, often only formed as marginal line. Caespituli amphigenous, punctiform to effuse, dark brown to blackish, later greyish by abundant conidiation. Mycelium internal; hyphae branched, septate, occasionally constricted at septa, 1.5–5 µm wide, subhyaline to brownish, thin-walled, smooth. Stromata variable, almost lacking, small to medium in size, subglobose to somewhat irregular, 10–30 µm diam, occasionally larger, to 80 µm diam, substomatal to immersed, olivaceous-brown to dark brown, cells subglobose to somewhat angular-irregular in outline, 2–6(− 11) µm diam, wall to 0.8 µm wide. Conidiophores in small to moderately large fascicles, occasionally in large fascicles, divergent to dense, arising from substomatal swollen hyphal cells or stromata, emerging through stomata or erumpent, erect, straight, subcylindrical or attenuated towards the tip, slightly to distinctly geniculate-sinuous, unbranched or rarely branched, 5–50(− 80) × 2–6 µm, 0–4-septate, longer ones plainly pluriseptate, subhyaline to pale or medium olivaceous, olivaceous-brown or brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous or visible as truncate tips or shoulders, always unthickened and not darkened. Conidia solitary, obclavate-cylindrical, straight to curved, 20–110(− 150) × 2–4.5(− 5) µm, 0–10(− 12)-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex acute to obtuse, base short to sometimes long obconically truncate, 0.8–2 µm wide, hila unthickened, not darkened.

[Holotype: USA: Illinois: on Ruellia ciliosa, A. B. Seymour]. Neotype (designated here, MycoBank, MBT202783): USA: Missouri: near Emma, on Ruellia ciliosa, Aug. 1889, C. H. Demetrio [Ellis & Everh., N. Amer. Fungi 2477] (BPI 435187). Isoneotypes: Ellis & Everh., N. Amer. Fungi 2477, e.g. DAOM, FH.

Host range and distribution: On Dyschoriste oblongifolia, Justicia (gendarussa, procumbens), Ruellia (ciliosa, prostrata, strepens, tuberosa), Acanthaceae, Asia (China; India, Uttar Pradesh, West Bengal; Japan, Thailand), North America (USA, Alabama, Florida, Iowa, Illinois, Mississippi, Missouri, Oklahoma), South America (Brazil, Venezuela).

var. dimorpha U. Braun & Urtiaga, Feddes Repert. 119: 489 (2008). (Fig. 28b)

Illustration: Braun & Urtiaga (2008: 490, fig. 4).

Description: Differs in vivo from var. consociata by the formation of superficial hyphae with solitary conidiophores (superficial hyphae emerging through stomata, branched, septate, 1.5–2.5(− 3) µm wide, subhyaline to pale olivaceous, thin-walled, smooth; solitary conidiophores arising from superficial hyphae, lateral, erect, straight, subcylindrical-conical to somewhat geniculate-sinuous, unbranched, 5–30 × 1.5–4 µm, 0–1(− 2)-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores often reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous); otherwise as in var. consociata.

Holotype: Venezuela: Lara State: Quíbor, on Justicia galapagana, Acanthaceae, Jan. 2007, R. Urtiaga (HAL 2190 F).

Host range and distribution: Only known from the type collection.

Notes: Pseudocercospora consociata, known from Asia, North and South America on hosts of various genera of the Acanthaceae, is probably taxonomically heterogeneous. This species is morphologically rather variable in all basic traits. However, a further splitting needs cultures and molecular sequence analyses based on material from all continents and host genera involved. Type material of this species, collected by Seymour before 1884, could not be traced. Records of Pseudocercospora consociata on Dicliptera chinensis from China (Guo & Hsieh 1995, Guo et al. 1998) are undoubtedly incorrect and belong to P. diclipterae. The two species are morphologically barely distinguishable.

Pseudocercospora diclipterae (A.K. Kar & M. Mandal) Deighton, Trans. Brit. Mycol. Soc. 88: 388 (1987). (Fig. 29)

Fig. 29
figure29

Pseudocercospora diclipterae (K(M) IMI 135116, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora diclipterae A.K. Kar & M. Mandal, Trans. Brit. Mycol. Soc. 53: 337 (1969).

Literature: Crous & Braun (2003: 159), Kamal (2010: 171).

Illustration: Kar & Mandal (1969: 338, fig. 1).

Description: Leaf spots amphigenous, subcircular, 1–6 mm diam, scattered, sometimes confluent, uniformly yellowish to dull yellowish brown, later olivaceous by abundant fructification. Caespituli amphigenous, mostly hypophyllous, punctiform to effuse, brown or deep olivaceous. Mycelium internal. Stromata lacking or small, 10–20 µm diam, subglobose or somewhat oblong, substomatal, brown. Conidiophores in small to moderately large fascicles, usually 2–15, divergent, arising from substomatal hyphae or stromata, through stomata, erect, straight to curved or geniculate-sinuous, unbranched, 10–55 × 3–5 µm, 0–5-septate, yellowish brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores sometimes reduced to conidiogenous cells, 10–20 µm long, conidiogenous loci inconspicuous to subdenticulate, unthickened, not darkened. Conidia solitary, obclavate-subcylindrical, straight to curved, 30–150 × 3–4 µm, 2–16-septate, pale yellowish brown, olivaceous, thin-walled, smooth, apex subacute or subobtuse, base short obconically truncate, 1.5–2 µm wide, hila unthickened, not darkened.

Holotype: India: West Bengal, Darjeeling, Sevak Road, on Dicliptera chinensis [roxburghiana], 11 Nov. 1967, M. Mandal (K(M) IMI 135116). Isotype: BPI 435712.

Host range and distribution: On Dicliptera chinensis, Acanthaceae, Asia (?China; India, West Bengal).

Note: Records of Pseudocercosora consociata on Dicliptera chinensis, from China belong possibly to P. diclipterae (see notes under P. consiciata).

Pseudocercospora ecbolii (A.K. Kar & M. Mandal) Deighton, Trans. Brit. Mycol. Soc. 88: 388 (1987). (Fig. 30)

Fig. 30
figure30

Pseudocercospora ecbolii (K(M) IMI 135117, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora ecbolii A.K. Kar & M. Mandal, Trans. Brit. Mycol. Soc. 53: 338 (1969).

Literature: Crous & Braun (2003: 168), Kamal (2010: 172).

Illustration: Kar & Mandal (1969: 339, fig. 2).

Description: Leaf spots amphigenous, formed as veinlimited, yellowish discolorations, 4–10 mm diam, scattered, finally deep olivaceous below by abundant fructification. Caespituli hypophyllous, effuse, olivaceous. Mycelium internal. Stromata lacking or small, substomatal, 5–25 µm diam, subglobose to oblong, dark olivaceous to olivaceous-brown. Conidiophores in small to moderately large fascicles, mostly 2–15, arising from stromata, through stomata, divergent to moderately dense, erect, straight, subcylindrical-conical to geniculate-sinuous, unbranched or occasionally 1–2 times branched, 15–65 × 3–5 µm, 0–6-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores sometimes reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci inconspicuous to subdenticulate, unthickened, not darkened. Conidia solitary, narrowly obclavate-cylindrical, straight to curved, 20–100 × 2.5–4 µm, 1–10-septate, pale olivaceous to very pale olivaceous-brown, thin-walled, smooth, apex subacute or subobtuse, base obconically truncate, 1–2 µm wide, hila neither thickened nor darkened.

Holotype: India: West Bengal: 24-Parganas, Rajarhat, on Ecbolium ligustrinum, Acanthaceae, 16 Jul. 1967, M. Mandal (K(M) IMI 135117). Isotype: BPI 436048.

Host range and distribution: Only known from the type collection.

Pseudocercospora justiciae (F.L. Tai) Y.L. Guo & X.J. Liu, Mycosystema 4: 103 (1991). (Fig. 31)

Fig. 31
figure31

Pseudocercospora justiciae (based on Guo & Hsieh 1995: 4, fig. 3). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Basionym: Cercospora justiciae F.L. Tai, Lloydia 11: 47 (1948).

Literature: Chupp (1954: 24), Guo & Hsieh (1995: 2), Guo et al. (1998: 11).

Illustrations: Tai (1948: 44, fig. 11), Guo & Hsieh (1995: 4, fig. 3), Guo et al. (1998: 13, fig. 3).

Description: Leaf spots lacking or only formed as yellowish to yellowish brown discolorations on the upper leaf surface, circular, 2–5 mm diam, grey to greyish brown below, margin indefinite. Caespituli hypophyllous, effuse, sooty. Mycelium internal and external, superficial; hyphae emerging through stomata, branched, septate, 2–3.5 µm wide, pale olivaceous to olivaceous-brown, thin-walled, smooth. Stromata not developed. Conidiophores in loose fascicles, 2–11, arising from internal hyphae, through stomata or solitary, arising from superficial hyphae, erect, subcylindrical, flexuous, curved-sinuous to somewhat geniculate-sinuous, unbranched or branched, 50–130 × 4–6.5 µm, 3–8-septate, olivaceous-brown to pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous, occasionally subdenticulate. Conidia solitary, cylindrical to obclavate-subcylindrical, straight to curved, 40–110 × 3.5–6.5 µm, 3–11-septate, pale olivaceous to olivaceous, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, about 2–2.5 µm wide, hila neither thickened nor darkened.

Holotype: China: Sichuan: Chengtu, on Justicia procumbens, 1943, Lee Ling 125 (HMAS 12126).

Host range and distribution: On Justicia (procumbens, Justicia sp.), Acanthaceae, Asia (China), ?North America (USA, Florida).

Notes: A record of P. justiciae on Justicia sp. from Florida, USA (Alfieri et al. 1984) is unclear and doubtful. Material could not be traced.

Pseudocercospora justiciicola P.N. Singh et al., Mycol. Res. 100: 1129 (1996). (Fig. 32)

Fig. 32
figure32

Pseudocercospora justiciicola (K(M) IMI 366368, holotype). A. Conidiophore fascicle. B. Solitary conidiophores arising from a superficial hypha. C. Conidiophore. D. Conidia. Bar = 10 µm.

Illustration: Singh et al. (1996: 1129, figs 15).

Description: Leaf spots lacking or almost so, lesions indistinct, occasionally formed as greyish discolorations. Caespituli hypophyllous, greyish, discrete to effuse. Mycelium internal and external; superficial hyphae branched, septate, subhyaline, 1–3 µm diam, thin-walled, smooth. Stromata developed, small, about 10–25 µm diam, subglobose to somewhat oblong, brown. Conidiophores in loose fascicles, arising from stromata or solitary arising from superfial hyphae, lateral, occasionally terminal, erect, straight to flexuous, somewhat geniculate-sinuous, unbranched, 20–50 × 1–5 µm, 2–6-septate, olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci unthickened, not darkened. Conidia solitary, obclavate, straight to curved, occasionally with short lateral branchlets, 35–120 × 1.5–5 µm, 4–14-septate, olivaceous, thin-walled, smooth, apex acute to subobtuse, base short obconically truncate, about 1–2 µm wide, hila unthickened, not darkened.

Holotype: Nepal: Kathmandu valley, on Justicia japonica [simplex], Acanthaceae, Nov.–Dec. 1993, Kamal (K(M) IMI 366368). Isotype: GPU 4004.

Host range and distribution: Only known from the type collection.

Notes: There is a second collection from Nepal that probably pertains to P. justiciicola. Verma & Kamal (1991) cited it as paratype material of P. rungiae [on “Ruellia prostrata” (on the original label as Justicia sp.), Nepal, Kathmandu Valley, Jan. 1986, R. K. Verma (GPU, KK 221, K(M) IMI 303480)]. See discussion under P. rungiae.

Pseudocercospora lepidagathidis U. Braun & Crous, Mycotaxon 92: 399 (2005). (Fig. 33)

Fig. 33
figure33

Pseudocercospora lepidagathidis (BPI 437637, holotype). A. Solitary conidiophores arising from superficial hyphae. B. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun & Crous (2005: 402, fig. 5).

Description: Leaf spots amphigenous, 1–5 mm diam, subcircular to somewhat irregular or even diffuse, brownish, dingy olivaceous-brown to blackish brown or with blackish purple tinge, margin indistinct. Caespituli hypophyllous, subeffuse, rather inconspicuous. Mycelium internal and external; superficial hyphae emerging through stomata, sparingly branched, 1–3 µm wide, septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth. Stromata lacking or only with very small substomatal hyphal aggregations, brown. Conidiophores in small, usually loose fascicles, arising from internal hyphae or stromatic hyphal aggregations, through stomata, or conidiophores solitary, arising from superficial hyphae, lateral or terminal, erect, straight to flexuous, geniculate-sinuous, unbranched or only rarely branched, 5–80 × 2.5–5.5 µm, 0–4-septate, pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–25 µm long, conidiogenous loci neither thickened nor darkened, occasionally subdenticulate. Conidia solitary, straight to curved, narrowly obclavate-cylindrical, 20–120 × 2–4 µm, 1–9-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: Puerto Rico: San Juan, on Lepidagathis alopecuroidea, Acanthaceae, 10 Feb. 1962, H. L. Rubin 16490 (BPI 437637).

Host range and distribution: Only known from the type collection.

Note: The material on Lepidagathis alopecuroidea was previous falsely referred to as Cercospora lepidagathidis (Stevenson 1975, Minter et al. 2001, Crous & Braun 2003), which is, however, a genuine species of Cercospora s. str.

Pseudocercospora odontonematis (Chupp) U. Braun & Crous, Mycosphaerella and Anam.: 296 (2003). (Fig. 34)

Fig. 34
figure34

Pseudocercospora odontonematis (CUP 40422, lectotype). A. Superficial hypha, B. Solitary conidiophores arising from superficial hyphae. C. Conidiophore fascicle. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Cercospora odontonematis Chupp, Monograph of Cercospora: 25 (1954); as “odontonemae”.

Literature: Chupp (1954: 25).

Description: Leaf spots amphigenous, circular to somewhat angular-irregular, 2–5 mm diam, brown, later with greyish brown to dingy grey centre and darker border. Caespituli hypophyllous, punctiform to effuse. Mycelium internal and external; superficial hyphae branched, septate, 1–3 µm wide, subhyaline or pale olivaceous, thin-walled, smooth; stromata substomatal, 10–40 µm diam, brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata or erumpent, also solitary, arising from superfial hyphae, lateral, rarely terminal, erect, straight, subcylindrical-conical to strongly geniculate-sinuous, unbranched, 10–40 × 2–4.5 µm, 0–2-septate, very pale or pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiophores recuced to conidiogenous cells or integrated, terminal, 10–25 µm long, conidiogenous loci inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, narrowly cylindrical to obclavate-cylindrical, 20–90 × 2–4 µm, (2−)3–8-septate, very pale olivaceous, thin-walled, apex obtuse to subacute, base more or less truncate to distinctly obconically truncate, 1.5–2 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202784): Mexico: Veracruz: Cordoba, on Odontonema callistachyum,

Acanthaceae, 25 Jul. 1932, O. A. Plunkett (CUP 40422). Isolectotype: CUP 60863.

Host range and distribution: Only known from the type collection.

Pseudocercospora rhinacanthi (Höhn.) Deighton, Mycol. Pap. 140: 152 (1976). (Fig. 35)

Fig. 35
figure35

Pseudocercospora rhinacanthi (BPI 440801, lectotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from a superficial hypha. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Cercospora rhinacanthi Höhn., Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl., Wien, 121: 414 (1912); as “rhynacanthr”.

Synonym: Cercosporina rhinacanthi (Höhn.) Sacc., Syll. Fung. 25: 917 (1931).

Literature: Chupp (1954: 25), Guo & Hsieh (1995: 357), Guo et al. (1998: 376), Crous & Braun (2003: 351).

Exsiccatae: Kabát & Bubák, Fungi Imp. Exs. 847.

Description: Leaf spots amphigenous, formed as diffuse yellowish to brown discolorations or diffuse brown spots with yellowish halo to circular or angular-irregular, 2–10 mm diam, brown or later with dingy grey centre, rather indistinct on dry leaves. Caespituli amphigenous, mainly hypophyllous. Mycelium internal and external; superficial hyphae branched, septate, 1–4 µm wide, subhyaline to olivaceous-brown, thin-walled, smooth. Stromata lacking or small, substomtal, 10–25 µm diam, olivaceous-brown to brown. Conidiophores in loose to moderately dense, small fascicles, usually 2–6, arising from stromata, through stomata, erect to decumbent, or solitary, arising from superficial hyphae, lateral, straight, subcylindrical to moderately geniculate-sinuous, unbranched or occasionally branched, 5–60(− 80) × 2–5 µm, 0–3-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiophores reduced to conidiogenous cells or integrated, terminal, about 10–25 µm long, conidiogenous loci inconspicuous or visible as truncate, subdenticulate tips or shoulders. Conidia solitary, narrowly obclavate-cylindrical, straight to somewhat curved, (25−)40–120(− 130) × 2–5 µm, indistinctly (2−)3–10(− 12)-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex more or less pointed, base usually long obconically truncate, 1–1.5 − m wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202785): Indonesia: Java: Bogor (Buitenzorg), Botanic Garden, on Rhinacanthus sp., 1907, F. v. Höhnel [Kabát & Bubák, Fungi Imp. Exs. 847] (BPI 440801). Isolectotypes: Kabát & Bubák, Fungi Imp. Exs. 847, e.g., K(M) IMI 89002 (slide), HBG, W.

Host range and distribution: On Rhinacanthus (nasutus [Justicia nasuta], Rhinacanthus sp.), Acanthaceae, Asia (Indonesia, Philippines, Thailand).

Notes: Chinese records of P. rhinacanthi on Justicia procumbens (Tai 1979, Guo & Hsieh 1995, Guo et al. 1998) are unclear, unproven and might belong to P. consociata. The two species are morphologically barely distinguishable.

Pseudocercospora rungiae R.K. Verma & Kamal, Indian Phytopathol. 44: 446 (1991). (Fig. 36)

Fig. 36
figure36

Pseudocercospora rungiae (K(M) IMI 299176, isotype). A. Conidiophore fascicle. B. Conidiophore (branched). C. Solitary conidiophores arising from a superficial hypha. D. Conidia. Bar = 10 µm.

Synonym: Cercospora rungiae M.S. Patil, Botanique (Nagpur) 8: 69 “1977” (1978) [holotype: India: Maharashtra: Kolhapur, on Rungia repens, 11 Dec. 1975, M. S. Patil (HCIO 32003)].

Literature: Crous & Braun (2003: 361), Kamal (2010: 216).

Illustrations: Patil (1978: 70, fig. 1), Verma & Kamal (1991: 444–445, figs 45).

Description: Leaf spots mainly hypophyllous, circular to angular-irregular or diffuse, 3–5 mm diam, greenish white, yellowish. Caespituli hypophyllous, scattered, rather inconspicuous. Mycelium internal; hyphae branched, septate, colourless, 1–2 µm wide, somewhat wider near stromata. Stromata lacking or poorly developed, substomatal, small, to about 25 µm diam. Conidiophores fasciculate, 2–10, divergent, arising from small stromata, through stomata, erect, straight, subcylindrical-conical to distinctly geniculate-sinuous, unbranched to branched, 10–75 × 2–5 µm, 0–3(− 6)-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiophores occasionally reduced to conidiogenous cells, but mostly integrated, terminal, 10–30 µm long, conidiogenous loci inconspicuous to subdenticulate, but always unbranched and not darkened. Conidia solitary, obclavate-subcylindrical, straight to curved, 30–145 × 2–4 µm, 1–10-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse or subacute, base short obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: India: Uttarakhand: Kath Godam, Nainital, on Rungia pectinata, Oct. 1985, R. K. Verma (GPU, KK 125). Isotype: K(M) IMI 299176.

Host range and distribution: On Rungia (pectinata [parviflora], repens), Acanthaceae, Asia (India, Maharashtra, Uttarakhand; Nepal).

Note: The identity of Cercospora rungiae and Pseudocercospora rungiae has been proven by Kamal (2010). A record of C. rungiae from Andhra Pradesh (Braun et al. 1992: 363) refers to a true Cercospora (see C. justiciicola) and not to this species. Verma & Kamal (1991) cited a collection on “Ruellia prostrata” (on the original label as Justicia sp.) as paratype material (Nepal: Kathmandu Valley, Jan. 1986, R. K. Verma (GPU, KK 221, K(M) IMI 303480). This collection, characterised by forming superficial hyphae with solitary conidiophores is excluded from P. rungiae. The identification of the host is uncertain, and the identity of this fungus is not quite clear, but it probably belongs to Pseudocercospora justiciicola.

Pseudocercospora thunbergiae (Boedijn) U. Braun & Sivapalan, Fungal Diversity 3: 21 (1999). (Fig. 37)

Fig. 37
figure37

Pseudocercospora thunbergiae (L 53879, holotype). A. Conidiophores and hyphae emerging through a stoma. B. superficial hyphae. C. Solitary conidiophore arising from a superficial hypha. D. Conidiophores. E. Conidia. Bar = 10 µm.

Basionym: Cercospora thunbergiae Boedijn, Nova Hedwigia 3: 411 (1961).

Literature: Crous & Braun (2003: 403), Kamal (2010: 225).

Illustrations: Boedijn (1961: tab. 108, fig. 1), Braun & Sivapalan (1999: 20, fig. 13).

Description: Leaf spots amphigenous, angular-irregular, often vein-limited, 1–10 mm diam, sometimes confluent and larger, dark brown, later greyish brown to dingy grey, margin indefinite. Caespituli hypophyllous, inconspicuous. Mycelium internal and external; superficial hyphae sparingly branched, 1–3 µm wide, septate, subhyaline to pale olivaceous, thin-walled, smooth. Stromata lacking or very small, only composed of a few swollen hyphal cells, 1.5–4 µm diam, olivaceous to brownish. Conidiophores solitary, arising from superficial hyphae, lateral, rarely terminal, occasionally emerging through stomata, solitary or in small, loose fascicle, erect, straight, subcylindrical-conical to somewhat geniculate-sinuous, unbranched, 5–30 × 1.5–3 µm, 0–1-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores aseptate, reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous, neither thickened nor darkened. Conidia solitary, narrowly obclavate-cylindrical, 20–95 × 1.5–4 µm, (0−)1–8-septate, subhyaline, pale yellowish green to very pale olivaceous, thin-walled, smooth, apex subacute, base obconically truncate, 1–2.5 µm wide, hila unthickened, not darkened.

Holotype: Indonesia: Jawa Barat: Bogor, botanical garden, Thunbergia alata, Apr. 1950, K. B. Boedijn (L 53879). Isotype: K(M) IMI 91586.

Host range and distribution: On Thunbergia (alata, erecta, grandiflora), Acanthaceae, Asia (Brunei; India, Uttar Pradesh; Indonesia).

Pseudocercospora thunbergiicola (J.M. Yen) Deight-on, Mycol. Pap. 140: 154 (1976). (Fig. 38)

Fig. 38
figure38

Pseudocercospora thunbergiicola (PC, holotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Cercospora thunbergiicola J.M. Yen, Rev. Mycol. 30: 195 (1965); as “thunbergicola”.

Literature: Yen & Lim (1980: 188), Crous & Braun (2003: 403).

Illustrations: Yen (1965: 197, fig. 12), Yen & Lim (1980: 258, fig. 68),

Description: Leaf spots amphigenous, circular to angular-irregular, 0.5–3 mm diam, centre pale, whitish, margin dark brown. Caespituli amphigenous, not very conspicuous. Mycelium internal and external; superficial hyphae emerging through stomata, sparingly branched, 2–3 µm wide, pale olivaceous-brown, thin-walled, smooth. Stromata almost lacking or small, 10–30 µm diam, substomatal, subglobose, brown. Conidiophores in small to moderately large fascicles, 2–22, arising from substomatal hyphae or stromata, emerging through stomata, divergent to moderately dense, or solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical or somewhat narrower towards the tip, to geniculate-sinuous, unbranched or occasionally once branched, 10–80 × 3–5 µm, 0–5-septate, pale olivaceous or olivaceous-brown, somewhat paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, conidiogenous loci inconspicuous or visible as truncate tips or shoulders. Conidia solitary, cylindrical or obclavate-cylindrical, straight to curved, about 60–175 × 3.5–5 µm, 4–11-septate, pale olivaceous-brown, thin-walled, smooth, apex subacute, base short to long obconically truncate, 2–2.5 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202786): Singapore: Botanic Garden, on Thunbergia alata, 20 Jun. 1964, S. H. Yen 60 (PC). Isolectotype: K(M) IMI 120991.

Host range and distribution: On Thunbergia alata, Acanthaceae, Asia (Singapore).

Semipseudocercospora

A single species.

Semipseudocercospora peristrophes-acuminatae (J.M. Yen) J.M. Yen, Mycotaxon 17: 363 (1983). (Fig. 39)

Fig. 39
figure39

Semipseudocercospora peristrophes-acuminatae (PC, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora peristrophes-acuminatae J.M. Yen, Rev. Mycol. 29: 230 (1964).

Literature: Yen & Lim (1980: 163), Crous & Braun (2003: 316).

Illustrations: Yen (1964: 233, fig. 10), Yen & Lim (1980: 219, fig. 22), Yen (1983: 362, fig. A–B).

Description: Leaf spots lacking or rather indistinct, forming diffuse dark brown discolorations. Caespituli hypophyllous, rarely amphigenous, effuse, brown. Mycelium internal. Stromata lacking. Conidiophores solitary or in small, divergent fascicles, arising from internal hyphae, through stomata, erect, straight to curved or somewhat flexuous, long, filiform, not or barely geniculate, unbranched, 140–260(− 300) × 5–7 µm, 3–15-septate throughout, medium brown, paler towards the tip, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, conidiogenous loci subconspicuous, 1.5–2 µm diam, in front view visible as small circle with darker rim and minute central pore, unthickened, but somewhat refractive or slightly darkened-refractive, loci somewhat bulging, denticle-like. Conidia solitary, short obclavate or clavate, ellipsoid, subcylindrical or obovoid, usually straight, 30–45 × 5.5–8.5 µm, pale olivaceous to olivaceous-brown, thin-walled, smooth, apex broadly rounded, base short obconically truncate, 1.5–2 µm wide, hila unthickened, but occasionally slightly refractive or darkened-refractive.

Lectotype (designated here, MycoBank, MBT202787): Singapore: Katong, on Peristrophe acuminata, Acanthaceae, S. H. Yen 20 (PC). Isolectotype: K(M) IMI 122324.

Host range and distribution: Only know from the type collection.

Notes: The generic affinity of Semipseudocercospora is quite unclear, and it is unknown if it is a true cercosporoid genus (Braun et al. 2013). Molecular sequence analyses are necessary to elucidate the phylogenetic position of this genus.

Actinidiaceae

Cercospora species on Actinidiaceae

A single species

Cercospora actinidiae X.J. Liu & Y.L. Guo, Acta Mycol. Sin. Suppl. 1: 353 “1986” (1987). (Fig. 40)

Fig. 40
figure40

Cercospora actinidiae (based on Guo et al. 2005: 23, fig. 6). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Guo et al. (2005: 23).

Illustrations: Liu & Guo (1987: 354, fig. 1), Guo et al. (2005: 23, fig. 6).

Description: Leafspots amphigenous, subcircular to irregular, 2–8 mm diam, centre greyish brown or yellowish brown, with narrow dark brown border. Caespituli amphigenous. Mycelium inernal. Stromata lacking or small, composed of a few swollen hyphal cells, brown. Conidiophores solitary or in small fascicles, 2–8, divergent, arising from internal hyphae or swollen hyphal cells, erect, straight, curved to geniculate, usually unbranched, about 60–280 × 4–4.5(− 5.5) µm, pluriseptate, medium brown to brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened, 2.2–2.6 µm diam. Conidia solitary, acicular, straight to curved, sometimes sigmoid, 40–160(− 220) × 2–3(− 4) µm, pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate or slightly attenuated, 1–2.5 µm wide, hila thickened and darkened.

Holotype: China: Jiangsu: Nanjing, on Actinidia sp., Actinidiaceae, 11 Sep. 1961, X. J. Liu & Q. M. Ma 249 (HMAS 50000).

Host range and distribution: Only known from the type collection.

Note: Belonging to the C. apii s. lat. complex.

Pseudocercospora species on Actinidiaceae

Key to Pseudocercospora species on Actinidiaceae

1 Conidia broadly obclavate-subcylindrical, 25–85(− 100) × 5–8.5 µm, base 2–2.5 µm wide; on Actinidia spp ………. P. actinidiae

Conidia narrowly obclavate-filiform, 15–110 × 1–4 µm, base 1–2 µm wide ………. P. hangzhouensis

Pseudocercospora actinidiae Deighton, Mycol. Pap. 140: 10 (1976). (Fig. 41)

Fig. 41
figure41

Pseudocercospora actinidiae (FH, holotype). A. Stroma. B. Conidiophore fascicles. C. Solitary conidiophores arising from superficial hyphae. D. Conidiophores. E. Conidia. Bar = 10 µm.

Literature: Liu & Guo (1987: 355), Hsieh & Goh (1990: 12–13), Guo & Hsieh (1995: 4–5), Guo et al. (1998: 15–16).

Illustrations: Deighton (1976: 11–12, figs 12), Liu & Guo (1987: 356, fig. 2), Hsieh & Goh (1990: 14, fig. 3), Guo & Hsieh (1995: 6, fig. 5), Guo et al. (1998: 15–16, fig. 5).

Exsiccatae: Reliquiae Farlowianae 829.

Description: Leaf spots at first lacking, later subcircular to angular-irregular, 1–10 mm diam, sometimes confluent and larger, brown to dark brown, later with paler centre, pale brown to greyish brown, margin indefinite. Caespituli amphigenous, mainly hypophyllous, finely punctiform to effuse, deep olivaceous, floccose. Mycelium internal and external; superficial hyphae emerging through stomata, branched, 1–3.5 µm wide, sometimes anastomosing, pale olivaceous to pale brown, thin-walled, smooth. Stromata lacking below to well-developed on the upper leaf surface, 10–50 µm diam, substomatal, subglobose, pale brown. Conidiophores in loose to dense fascicles, to 100 conidiophores or even more, arising from stromata, through stomata, above all when epiphyllous, and solitary, arising from superficial hyphae, lateral, occasionally terminal, differentiation between hypophyllously formed erect to decumbent branched conidiophores and superficial hyphae difficult, fertile threads to 700 µm long, simple or branched, individual solitary conidiophores formed as lateral branchets 2–45 µm long, 3–6.5 µm wide, aseptate to pluriseptate, pale to medium olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous to subdenticulate, 1.5–2 µm diam, unthickened, not darkened. Conidia solitary, obclavate-subcylindrical, straight to curved or somewhat sigmoid, 25–85(− 100) × 5–8.5 µm, 3–9-septate, pale to medium olivaceous, thin-walled, smooth, apex obtuse, base short or rarely long obconically truncate, about 2–2.5 µm wide, hila unthickened, not darkened.

Holotype: China: Guizhou (Kweichow): Fan Ching Shan, on Actinidia callosa, 21 Sep. 1931, S. Y. Cheo 570 (FH). Isotypes: BPI 437420, 876607; HMAS 12125 and Reliquiae Farlowianae 829.

Host range and distribution: On Actinidia (callosa, chinensis, Actinidia sp.), Actinidiaceae, Asia (China, Anhui, Fujian, Guizhou; Korea, Japan, Taiwan).

Pseudocercospora hangzhouensis X.J. Liu & Y.L. Guo, Acta Mycol. Sin. Suppl. 1: 357 “1986” (1987). (Fig. 42)

Fig. 42
figure42

Pseudocercospora hangzhouensis (based on Guo & Hsieh 1995: 7, fig. 6). A. Conidiophore fascicle. B. Solitary conidiophores arising from superficial hyphae. C. Conidia. Bar = 10 µm. U. Braun del.

Synonym: Pseudocercospora actinidiicola Goh & W.H. Hsieh, Bot. Bull. Acad. Sin. (Taipei) 30: 122 (1989) [holotype: Taiwan: Taichung, NCHU Campus, on Actinidia chinensis, 14 Aug. 1984, T. K. Goh (NCHUPP-49); isotype: K(M) IMI 312075].

Literature: Hsieh & Goh (1990: 12–13), Guo & Hsieh (1995: 5–6), Guo et al. (1998: 16–17).

Illustrations: Liu & Guo (1987: 357, fig. 3), Hsieh & Goh (1990: 15, fig. 4), Guo & Hsieh (1995: 7, fig. 6), Guo et al. (1998: 17, fig. 6).

Description: Leaf spots amphigenous, subcircular to angular-irregular and vein-limited, 1–18 mm diam, brown to dark brown, yellowish brown below, later centre greyish white with dark margin, sometimes with yellowish brown halo. Caespituli amphigenous. Mycelium internal and external; superficial hyphae branched, 1.5–3 µm wide, subhyaline to olivaceous, thin-walled, smooth. Stromata 10–70 µm diam, subglobose to oblong, mainly epiphyllous, brown. Conidiophores in dense fascicles, arising from stromata, through stomata or erumpent, or solitary, arising from superficial hyphae, erect, straight to curved, subcylindrical to geniculate-sinuous, unbranched, 2–40 × 1–3.5(− 4.5) µm, 0–3-septate, subhyaline to pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 2–20 µm long, conidiogenous loci inconspicuous. Conidia solitary, narrowly obclavate-filiform, 15–110 × 1–4 µm, 2–11-septate, hyaline to pale olivaceous, thin-walled, smooth, apex subacute, base subtruncate to short obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: China: Zhejiang: Hangzhou, on Actinidia arguta, 28 Sep. 1961, X.-J. Liu & Q.-M. Ma 671 (HMAS 50001).

Host range and distribution: On Actinidia (arguta, chinensis, Actinidia sp.), Actinidiaceae, Asia (China, Anhui, Zhejiang; Japan, Korea, Taiwan).

Doubtful, excluded and insufficiently known species

Pseudocercospora clematoclethrae X.J. Liu & Y.L. Guo, Mycosystema 5: 100 (1992).

Notes: Guo & Liu (1992) described P. clematoclethrae from China on “Clematoclethra fabri” [C. scandens subsp. actinidioides] (Actinidiaceae), but later Guo & Hsieh (1995) and Guo et al. (1998) changed the identification of the host to Clethra sp. (Clethraceae).

Adoxaceae

Cercospora

Tabular key to Cercospora species on Adoxaceae according to host genera
  • Sambucus

    A single species ……… C. sambucicola

  • Viburnum

    A single species ………. C. viburnicola

Cercospora species on Adoxaceae

Cercospora sambucicola Y.L. Guo, nom. nov. MycoBank MB814566 (Fig. 43)

Fig. 43
figure43

Cercospora sambucicola (HMAS 77346, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora sambuci Y.L. Guo & Y. Jiang, Mycotaxon 74: 262 (2000), nom. illeg. (Art. 53.1), non C. sambuci F. Stevens & C.J. King, 1927.

Literature: Guo et al. (2005: 55–56).

Illustrations: Guo & Jiang (2000: 263, fig. 3), Guo et al. (2005: 56, fig. 32).

Description: Leaf spots amphigenous, circular or subcircular, 2–8 mm diam, pale yellowish brown to brown, later with greyish white centre on the upper leaf surface, surrounded by a greyish black marginal line, pale yellowish brown to pale greyish brown below. Caespituli amphigenous. Mycelium internal. Stromata lacking or small, only formed as aggregations of a few swollen hyphal cells, globose, 10–25 µm diam, substomatal or intraepidermal, brown, cells 2–8 µm diam, wall slightly thickened. Conidiophores solitary or in small, loose fascicles, 2–13, arising from internal hyphae or hyphal aggregations, through stomata or erumpent, erect, straight, subcylindrical to geniculate-sinuous, unbranched, width irregular, 42.5–140(− 400) × 3.5–7 µm, to 8 µm wide at the base, 1–6(− 13)-septate, medium brown, paler and narrower towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened, 2–3.5(− 4) µm diam. Conidia solitary, acicular, straight to somewhat curved or slightly sigmoid, 37.5–175 × 3–5 µm, 5- to pluriseptate, hyaline, thin-walled, smooth, apex pointed or subobtuse, base truncate, 2–3.5 µm wide, hila thickened and darkened.

Holotype: China: Jilin Province: Yongji, on Sambucus williamsii, Adoxaceae, Sep. 1962, J. K. Bai (HMAS 77346).

Host range and distribution: On Sambucus (?canadensis, javanica, williamsii [buergeriana]), Adoxaceae, Asia (China, Jilin, Zhejiang), ?North America (USA, Alabama).

Notes: This species belongs to the Cercospora apii s. lat. complex. Type material of Passalora catenospora contains a second cercosporoid hyphomycete which is morphologically barely distinguishable from C. sambucicola (conidiophores in small fascicles, to 140 µm long and 4–7 µm wide; conidia acicular, hyaline, 3–4.5 µm wide). The conspecifity of Chinese collections and material from the USA remains to be proven via phylogenetic analyses.

Cercospora viburnicola W.W. Ray, Mycologia 33: 174 (1941). (Fig. 44)

Fig. 44
figure44

Cercospora viburnicola (CUP 29236, holotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 106), Crous & Braun (2003: 421).

Illustration: Guo et al. (2005: 57, fig. 33).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 2–12 mm diam, different shades of brown, pale to medium brown, reddish brown to greyish brown. Caespituli amphigenous, fine, dark. Mycelium internal. Stromata lacking or small, about 10–25 µm diam, brown. Conidiophores in small to moderately large fascicles, mostly dense, arising from internal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 25–175 × 4–6 µm, septate throughout, uniformly pale olivaceous-brown or somewhat paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, about 15–35 µm long, conidiogenous loci conspicuous, thickened and darkened, 2–3 µm diam. Conidia solitary, acicular, shorter conidia narrowly obclavate, straight to curved, 20–90 × 2–4 µm, pluriseptate, hyaline, thin-walled, smooth, apex acute or subacute, base truncate or somewhat obconically truncate in obclavate conidia, 2–2.5 µm wide, hila thickened and darkened.

Holotype: USA: Oklahoma: Payne County, Stillwater, college garden, on Viburnum opulus, 18 Aug. 1939, W. W. Ray 286 (CUP 29236).

Host range and distribution: On Viburnum (carlesii, odoratissimum, opulus, plicatum var. tomentosum [tomentosum], suspensum, Viburnum spp.), Adoxaceae, ?Asia (China, Jiangsu, Shaanxi, Sichuan), USA (Florida, Oklahoma, Wisconsin).

Notes: Cercospora viburnicola W.W. Ray is morphologically part of the Cercospora apii s. lat. complex. Reords of this species on Viburnum cylindricum from China are incorrect and based on a confusion with C. viburnicola F.L. Tai. Authentic material collected at the type locality (21 Aug. 1942, on Viburnum tomentosum) is deposited in several herbaria (BPI 442264–442266, CUP-C.H.OK-0032, ILL 29523, NY 945736). A record of C. viburnicola from Poland on Viburnum opulus (Świderska-Burek & Mułenko 2014) is undoubtedly based on confusion and misidentification. The described and illustrated obclavate conidia with obconically truncate base, 30–90(− 205) × 3–5(− 5.5) µm, are not consistent with C. viburnicola. The collection concerned has been re-examined and turned out to belong to Pseudocercospora opuli which is common on Viburnum opulus and also known from Poland. Chinese collections on Viburnum buddleifolium and Viburnum sp. (Guo 1996: 92; Guo & Jiang 2000: 264; Guo et al. 2005: 56–57, fig. 33), referred to as C. viburnicola, represent typical C. apii s. lat., but these specimens are probably not conspecific with the latter species. The conidiophores are formed in small, divergent fascicles, to about 450 µm long, and the conidia are acicular, about 35–385 µm long. Without any molecular data based on Asian and North Americal samples, the Chinese collections are currently better classified as Cercospora apii s. lat.

Doubtful, excluded and insufficiently known species

Cercospora adoxae Roum., Fungi Sel. Gall. Exs., Cent. XIX, no. 1871, Toulouse 1882, nom. nud. (Art. 38.1).

Literature: Chupp (1954: 98), Crous & Braun (2003: 46).

Type: France: on Adoxa moschatellina, Roum., Fungi Sel. Gall. Exs. 1871 (e.g., B, HBG, PC).

Host range and distribution: On Adoxa moschatellina, Adoxaceae, Europe (France).

Notes: Although listed in literature, this species was never described (Chupp 1954). Numerous duplicates of the original material have been examined, but all were without any trace of fructification. The name C. adoxae might refer to Ramularia adoxae (Rabenh.) P. Karst.

Cercospora prolificans Ellis & Holw., Bull. Lab. Nat. Hist. Iowa State Univ. II, 3: 42 (1896).

Synonym: Cercosporella prolificans (Ellis & Holw.) Sacc., Syll. Fung. 15: 84 (1901).

Literature: Saccardo (1895: 606), Chupp (1954: 104), Braun (1995: 80).

Illustration: Braun (1995: 79, fig. 75).

Exsiccatae: Calif. Fungi 403.

Lectotype (designated by Braun 1995: 80): USA: California: San Bernadino Valley, San Bernadino County, on Sambucus cerulea, Aug. 1893, Parish 2735 (NY 234139). Isolectotypes: BPI 439986, NY 234138.

Host range and distribution: On Sambucus (cerulea, glauca, melanocarpa), Adoxaceae, North America (USA, Alaska, California, Minnesota, Missouri, New Mexico, Oregon).

Notes: Cercospora prolificans has colourless conidiophores and conidia as well as cercosporelloid conidiogenous loci and belongs to Cercosporella (Braun 1995). This species was recorded and described from Bulgaria on Sambucus ebulus (Bakalova & Borisova 2010: 49–50). This record is, however, doubtful and probably wrong. Conidia were described to be 2–4 µm wide, which is in contradiction with the much broader conidia (3–8 µm wide) of true C. prolificans.

Passalora

Key to Passalora species on Adoxaceae
  1. 1

    Colonies in pale reddish brown to fuliginous patches; mycelium internal and external, superficial; conidiophores fasciculate as well as solitary, arising from superficial hyphae; conidia subhyaline to reddish brown; on Sambucus spp. ………. P. lateritia

    Colonies (caespituli) without reddish tinge; mycelium internal, superficial hyphae lacking in vivo; conidia not reddish brown ………. 2

  2. 2(1)

    Conidiophores short, 5–25 × 2–4 µm, 0–1-septate; conidia 8–45 × 1.5–4 µm, 0–4-septate; on Viburnum spp. ………. P. viburni

    Conidiophores much larger, 15–80 × 4–6 µm, 0–4-septate; conidia much longer and somewhat broader, 20–100(− 125) × 3–5.5 µm, 1–8-septate; on Sambucus spp. ………. P. catenospora

Passalora catenospora (G.F. Atk.) U. Braun & Crous, Mycosphaerella and Anam.: 112 (2003). (Fig. 45)

Fig. 45
figure45

Passalora catenospora (CUP-A-002255a, lectotype). A. Conidiophore fascicles. B. Conidiophore. C. Conidia. Bar = 10 µm.

Basionym: Cercospora catenospora G.F. Atk., J. Elisha Mitchell Sci. Soc. 8: 66 “1891” (1892).

Synonyms: Phaeoramularia catenospora (G.F. Atk.) Deighton, More Dematiaceous Hyphomycetes: 317 (1976).

Cercospora affinis G. Winter, in herb. [USA: Pennsylvania, on Sambucus canadensis, ex herb. G. Winter (B 700016202)].

Literature: Saccardo (1892: 645), Chupp (1954: 100).

Illustration: Ellis (1976: 318, fig. 240 A).

Exsiccatae: Barthol., Fungi Columb. 2309.

Description: Leaf spots indistinct. Caespituli forming effuse, irregular patches, hypophyllous, ochraceous, dark olivaceous to brownish. Mycelium internal. Stromata lacking, only formed as small aggregations of swollen hyphal cells or small, 10–25 µm diam, substomatal, yellowish to ochraceous-brown. Conidiophores in small to moderately large fascicles, usually 3–12, arising from swollen hyphal cells or small stromata, emerging through stomata, erect, straight, subcylindrical or attenuated towards the tip, not or only slightly geniculate, somewhat curved-sinuous, unbranched, 15–80 × 4–6 µm, rarely longer, usually 0–4-septate, subhyaline, pale ochraceous to golden brown or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, (1−)1.5–2 µm wide. Conidia solitary to catenate, in simple or branched chains, cylindrical, subcylindrical, occasionally almost obclavate or broadly fusiform, straight to curved or slightly sinuous, 20–100(− 125) × 3–5.5 µm, 1–8-septate, subhyaline, pale olivaceous to brownish, thin-walled, smooth, apex obtuse, rounded (in primary conidia) or short conically truncate in catenate conidia (with a single or two hila), base short obconically truncate, (1−)1.5–2 µm wide, hila somewhat thickened and darkened.

Lectotype (designated here, MycoBank, MBT202788): USA: Alabama: Lee County, Auburn, on Sambucus canadensis, 11 Oct. 1891, G. F. Atkinson (CUP-A-002255a). Isolectotype: CUP-A-002255b.

Host range and distribution: on Sambucus (cerulea, canadensis, intermedia, Sambucus spp.), Adoxaceae, Central America (Dominican Republic), North America (Canada; USA, Alabama, Florida, Kansas, Mississippi, North Carolina, Oklahoma, Pennsylvania, Texas), West Indies (Haiti).

Note: The lectotype is a collection from October 1891. Other syntypes are from August 1891 (CUP-A-002245#1–3(AL)). Records of this species from Taiwan on Sambucus javanica [formosana] (Tai 1979) are incorrect and refer to Pseudocercospora ebulicola (Cercospora ebulicola) which was erroneously reduced to synonymy with C. catenospora in Chupp (1954). Type material of C. catenospora contains a second cercosporoid hyphomycete which might cause confusion. The second fungus belongs to the Cercospora apii s. lat. complex and is morphologically barely distinguishable from the Chinese Cercospora sambucicola. This species is readily distinguishable from P. catenospora by its much darker conidiophores with larger conidiogenous loci, 2–3 µm diam, as well as hyaline, acicular conidia formed singly.

Passalora lateritia (Ellis & Halst.) U. Braun & Crous, Mycosphaerella and Anam.: 244 (2003). (Fig. 46)

Fig. 46
figure46

Passalora lateritia (NY 838232, lectotype). A. Superficial hypha. B. Conidiophore fascicles. C. Conidiophore. D. Solitary conidiophores arising from superficial hyphae. E. Conidia. Bar = 10 µm.

Basionym: Cercospora lateritia Ellis & Halst., J. Mycol. 4: 7 (1888).

Literature: Saccardo (1892: 646), Chupp (1954: 102).

Exsiccatae: Ellis & Everh., N. Amer. Fungi 1994. Kellerman & Swingle, Kansas Fungi 6.

Description: Leafspots lacking or indefinite. Colonies hypophyllous, effuse, forming reddish, pale reddish brown to ferruginous patches. Mycelium internal and external; superficial hyphae lacking or present, emerging through stomata, occasionally climbing leaf hairs, rarely anastomosing, unbranched or sparingly branched, 1–7 µm wide, septate, concolorous with conidiophores or pale, thin-walled, smooth. Stromata lacking or formed as stromatic aggregations of swollen hyphal cells, substomatal, 10–50 µm diam, reddish brown. Conidiophores in small to moderately large fascicles, divergent to moderately dense, arising from substomatal hyphae or stromata, through stomata or conidiophores solitary, arising from superficial hyphae, lateral or terminal, 5–75 × 3–8 µm, 0–4-septate, subhyaline to pale reddish brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–30 µm long, conidiogenous loci conspicuous, thickened and darkened, about 1–2 µm diam. Conidia solitary to catenate, usually in simple chains, cylindrical or subcylindrical or somewhat obclavate-cylindrical, straight to slightly curved, 15–70(− 80) × 3–7 µm, (0−)1–6-septate, occasionally constricted at the septa, subhyaline to pale reddish brown, thin-walled, smooth, apex obtuse, rounded in solitary conidia, short conically truncate in catenate conidia, base obconically truncate, (1−)1.5–2(− 2.5) µm wide, hila somewhat thickened and darkened.

Lectotype (designated by Crous & Braun 2003: 244): USA: Iowa: Story County, Ames, on Sambucus pubens, Sep. 1887, B. D. Halsted (NY 838232). Isolectotypes: BPI 437793, CUP-040123, NY 838230, 838233; Ellis & Everh., N. Amer. Fungi 1994, e.g., BPI 437794, MICH 15321.

Host range and distribution: On Sambucus (canadensis, nigra, pubens, racemosa), Adoxaceae, North America (USA, Iowa, Kansas).

Notes: This is a typical mycovellosielloid member of Passalora, with fasciculate and solitary conidiophores arising from superficial hyphae, thickened, darkened conidiogenous loci and obclavate-cylindrical, pigmented conidia. This species is very characteristic by its effuse, reddish to reddish brown colonies and reddish tinge of conidiophores and conidia.

Passalora viburni (Ellis & Everh.) U. Braun & Crous, Mycosphaerella and Anam.: 474 (2003). (Fig. 47)

Fig. 47
figure47

Passalora viburni (NY 830558, lectotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Ramularia viburni Ellis & Everh., J. Mycol. 5: 69 (1889).

Synonym: Phaeoramularia viburni (Ellis & Everh.) U. Braun, Mycotaxon 48: 293 (1993).

Literature: Saccardo (1892: 554), Braun (1998: 387).

Illustrations: Braun (1993: 292, fig. 21), Braun (1998: 388, fig. 647).

Description: Leaf spots subcircular to angular-irregular, sometimes vein-limited, 2–10 mm diam, greyish green to brown, later centre greyish white, with darker margin. Caespituli amphigenous, punctiform, greyish white to brownish. Mycelium internal. Stromata intraepidermal, occasionally substomatal, 10–50 µm diam, occasionally confluent and larger, pale yellowish brown to medium brown, composed of swollen hyphal cells, 2–6 µm diam. Conidiophores in small to rather large fascicles, arising from stromata, erumpent, occasionally emerging through stomata, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 5–25 × 2–4 µm, 0–1-septate, hyaline, yellowish, greenish to pale olivaceous or yellowish brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores mostly reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci conspicuous, minute, 1–1.5 µm diam, barely or very slightly thickened, somewhat darkened-refractive. Conidia solitary or catenate, in simple or rarely branched chains, shape and size variable, narrowly ellipsoid-ovoid, fusiform, subcylindrical, straight to slightly curved, 8–45 × 1.5–4 µm, 0–4-septate, hyaline to pale yellowish or greenish, smooth to faintly verruculose, thin-walled, ends attenuated, short obconically truncate when in chains, about 1 µm wide, hila minute, very slightly thickened and darkened-refractive.

Lectotype (designated by Braun 1993): USA: Wisconsin: Racine, on Viburnum lentago, 17 Jun. 1888, J. J. Davis (NY 830558). Isolectotype: NY 830559, WIS.

Host range and distribution: On Viburnum (lentago, prunifolium), Adoxaceae, North America (USA, Tennessee, Wisconsin).

Notes: The generic affinity of this species, which is intermediate between Passalora and Ramularia, is not quite clear. The general habit of this species is rarther ramularioid, but the stromata and conidiophores are pigmented. Results of molecular sequence analyses would be helpful to elucidate the true generic affinity.

Pseudocercospora

Key to Pseudocercospora species on Adoxaceae
  1. 1

    Stromata lacking; superficial hyphae developed, but without solitary conidiophores; conidiophores fasciculate, 50–190 µm long, pluriseptate; on Sambucus javanica, Asia ………. P. ebulicola

    Stromata developed, 10–100 µm diam; superficial hyphae lacking or, if present, with conidiophores which are much shorter, 5–35 µm, and only 0–1-septate ………. 2

  2. 2(1)

    Superficial hyphae with solitary conidiophores developed, 5–35 µm long, 0–1-septate, geniculate, i.e. proliferation sympodial; on Viburnum spp ………. P. tinea

    Superficial hyphae with solitary conidiophores lacking; conidiophores longer, 10–200 µm, aseptate to pluriseptate (if short and mostly aseptate, then non-geniculate, proliferation percurrent, with annellations ………. 3

  3. 3(2)

    Conidiophores subcylindrical, non-geniculate, proliferation percurrent, with fine annellations, rather short, 5–30(− 50) × 2–5(− 6) µm, 0(− 2)-septate; conidia cylindrical-filiform or occasionally somewhat obclavate; on Viburnum spp ………. P. viburnigena

    Conidiophores at least partly geniculate-sinuous, proliferation sympodial, without annellations or occasionally with a single enteroblastic rejuvenation, much longer, 10–200(− 300) µm, 0–3-septate or pluriseptate throughout; conidia obclavate-cylindrical to somewhat fusiform ………. 4

  4. 4(3)

    Conidia short, 15–40 × 3–7 µm, 1–5-septate, pigmented; conidiophores in divergent fascicles, walls thickened, to 1 µm, medium to dark brown, occasionally with a single enteroblastic rejuvenation leaving a coarse annellation; on Viburnum nudum, North America ………. P. viburni-nudi

    Conidia longer, 25–150 µm, 1–12(µ15)-septate; on other hosts ………. 5

  5. 5(4)

    Conidia broadly obclavate or obclavate-cylindrical, 25–125 × 4.5–8 µm, 2–10-septate, usually pale to medium olivaceous or olivaceous-brown, wall thin to somewhat thickened, to 0.8 µm; on Viburnum spp., North America ………. P. viburnicola

    Conidia narrower, 2–5 µm, if wider [2–6.5(− 7) µm] conidia consistently pale, usually hyaline or subhyaline and/or thin-walled, or conidia at least parly catenate or disarticulating ………. 6

  6. 6(5)

    Conidiophores relatively short, usually 10–80 µm, average < 60 µm ………. 7

    Conidiophores longer, about 50–155 µm, average > 60 µm ………. 9

  7. 7(6)

    Conidia hyaline or subhyaline, finally sometime pale olivaceous, solitary or in short chains or disarticulating, conidiogenous loci inconspicuous to subconspicuous, i.e. unthickened, but somewhat refractive or even slightly darkened-refractive; on Viburnum spp. (Viburnum sect. Opulus), mainly V. opulus s. lat. (including subsp. calvescens and trilobum) ………. P. opuli

    Conidia formed singly ………. 8

  8. 8(7)

    Conidiogenous loci inconspicuous to subconspicuous, i.e. unthickened but somewhat refractive or even darkened-refractive; on Sambucus spp. ………. P. depazeoides

    Conidiogenous loci inconspicuous (conidiophores usually in small fascicles, 2–15, divergent, North American collections = var. varia; conidiophores in larger, sense fascicles, to 50 per fascicle, Asian collections on Viburnum sargentii = var. viburni-sargentii); on Viburnum spp. ………. P. varia

  9. 9(6)

    Conidiogenous loci inconspicuous to subconspicuous, i.e. unthickened but somewhat refractive or even darkened-refractive; conidiophores fascicluate; on Sambucus spp. ………. P. depazeoides

    Conidiogenous loci consistently inconspicuous; conidiophores fasciculate to coremioid; on Viburnum spp. ……… 10

  10. 10(9)

    Conidia cylindrical-filiform, 50–130 × 2.5–4 µm, 3–12-septate; on Viburnum erosum ………. P. viburni-erosi

    Conidia obclavate-cylindrical, 25–85 µm long, 2–7-septate ………. 11

  11. 11(10)

    Conidia 2.5–4 µm wide, 2–5-septate; on Viburnum cylindricum, China ………. P. viburni-cylindrici

    Conidia wider, 3–6.5 µm, 3–7-septate; on Viburnum sp., India ………. P. caprifoliacearum

Tabular key to Pseudocercospora species on Adoxaceae according to host genera

Sambucus

1 Stromata lacking; superficial hyphae developed; conidiophores loosely fasciculate, simple to branched; on Sambucus javanica, Asia ………. P. ebulicola

Stromata developed, 10–100 µm diam; superficial hyphae lacking; conidiophores mostly in denser fascicles, unbranched; on Sambucus spp., northern hemisphere ………. P. depazeoides

Viburnum
  1. 1

    Superficial hyphae with solitary conidiophores developed, 5–35 µm long, 0–1-septate, geniculate, i.e. proliferation sympodial ………. P. tinea

    Superficial hyphae with solitary conidiophores lacking; conidiophores longer, 10–200 µm, aseptate to pluriseptate (if short and mostly aseptate, then non-geniculate, proliferation percurrent, with annellations ………. 2

  2. 2(1)

    Conidiophores subcylindrical, non-geniculate, proliferation percurrent, with fine annellations, rather short, 5–30(− 50) × 2–5(− 6) µm, 0(− 2)-septate; conidia cylindrical-filiform or occasionally somewhat obclavate ………. P. viburnigena

    Conidiophores at least partly geniculate-sinuous, proliferation sympodial, without annellations or only with a single enteroblastic rejuvenation, much longer, 10–200(− 300) µm, 0–3-septate or pluriseptate throughout; conidia obclavate-cylindrical to somewhat fusiform ………. 3

  3. 3(2)

    Conidia broadly obclavate or obclavate-cylindrical, 25–125 × 4.5–8 µm, 2–10-septate, usually pale to medium olivaceous or olivaceous-brown, wall thin to somewhat thickened, to 0.8 µm; on Vibrunum spp., North America ………. P. viburnicola

    Conidia narrower, 2–5 µm, if wider [2–6(− 7) µm] conidia consistently pale, usually hyaline or subhyaline and thin-walled ………. 4

  4. 4(3)

    Conidiophores relatively short, usually 10–80 µm, average < 60 µm ………. 5

    Conidiophores longer, about 50–155 µm, average > 60 µm ………. 7

  5. 5(4)

    Conidia short, 15–40 × 3–7 µm, 1–4(− 5)-septate, pigmented; conidiophores in divergent fascicles, walls thickened, to 1 µm, medium to dark brown, occasionally with a single enteroblastic rejuvenation leaving a coarse annellation; on Viburnum nudum, North America ………. P. viburni-nudi

    Conidia longer, 30–150 µm, 1–12(− 15)-septate; on other hosts ………. 6

  6. 6(5)

    Conidia solitary, in short chains or disarticulating, conidiogenous loci inconspicuous to subconspicuous, i.e. unthickened, but somewhat refractive or even slightly darkened-refractive; on Viburnum spp. (Viburnum sect. Opulus), mainly V. opulus s. lat. (including subsp. calvescens and trilobum) ……….P. opuli

    Conidia solitary; conidiogenous loci inconspicuous, not darkened-refractive (conidiophores usually in small fascicles, 2–15, divergent, North American collections = var. varia; conidiophores in larger, sense fascicles, to 50 per fascicle, Asian collections on Viburnum sargentii = var. viburni-sargentii); on Viburnum spp ………. P. varia

  7. 7(4)

    Conidia cylindrical-filiform, 50–130 × 2.5–4 µm, 3–12-septate; on Viburnum erosum ………. P. viburni-erosi

    Conidia obclavate-cylindrical, 25–85 µm long, 2–7-septate ………. 8

  8. 8(7)

    Conidia 2.5–4 µm wide, 2–5-septate; on Viburnum cylindricum, China ………. P. viburni-cylindrici

    Conidia wider, 3–6.5 µm, 3–7-septate; on Viburnum sp., India ………. P. caprifoliacearum

Pseudocercospora caprifoliacearum (C. Gupta et al.) Kamal, Cercosporoid Fungi of India: 159 (2010). (Fig. 48)

Fig. 48
figure48

Pseudocercospora caprifoliacearum (K(M) IMI 294111, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Phaeoisariopsis caprifoliacearum C. Gupta et al., Perspectives in Mycological Research, Prof. G.P. Agarwal Festschrift Volume 1: 9 (1987).

Synonym: Pseudocercospora khasiana B.K. Gupta & Kamal, Perspectives in Mycological Research, Prof. G.P. Agarwal Festschrift Volume 1: 25 (1987) [holotype: India: Meghalaya, Shillong, on Viburnum sp., B. K. Gupta KB 58 (K(M) IMI 274850)].

Literature: Braun (1992: 219), Kamal (2010: 230).

Illustrations: Gupta et al. (1987: 16, fig. 2), Gupta & Kamal (1987: 33, fig. 4).

Description: Leaf spots amphigenous, subcircular to angular-irregular, 2–20 mm diam, brown, olivaceous-brown, later greyish brown to dingy grey, margin indefinite or with narrow to broad darker border or halo. Caespituli amphigenous, scattered, finely punctiform, brown to blackish. Mycelium immersed, composed of subhyaline to pale brown, septate, thin-walled, smooth hyphae. Stromata about 10–50 µm diam, dark brown, substomatal, cells 2–6 µm diam. Conidiophores in loose to dense, almost coremoid fascicles, 7–18, arising from stromata, though stomata, erect, when coremioid then dense below and looser towards the apex, straight to flexuous, unbranched or only rarely branched, subcylindrical-filiform, width uniform throughout or tips somewhat swollen, geniculate-sinuous in the upper half, about 60–155 µm long and 2.5–4.5 µm wide, apex sometime to 6 µm wide, pluriseptate throughout, olivaceous-brown, wall thin-walled to somewhat thickened, to 1 µm, at the very base occasionally to 1.3 µm, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, sympodially proliferating, with several conidiogenous loci, inconspicuous or visible as truncate tips or shoulders, subdenticulate, but not thickened. Conidia solitary, obclavate-cylindrical, broadly fusiform, straight to slightly curved, about 30–85 × 3–6.5 µm, 1–7-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse, base obconically truncate, 1.5–2.5 µm wide, hila unthickened, not darkened.

Holotype: India: Uttar Pradesh: Gorakhpur, on Viburnum sp., Adoxaceae, Jan. 1985, C. Gupta (K(M) IMI 294111). Isotype: GPU, KC 62.

Host range and distribution: On Viburnum spp., Adoxaceae, Asia (India, Meghalaya, Uttarakhand, Uttar Pradesh).

Notes: Pseudocercospora caprifoliacearum and P. khasiana, both described from India on Viburnum sp., are morphologically barely distinguishable and undoubtedly conspecific. The two species have been simultaneously published in a single book. It is proposed to give precedence to Phaeoisariopsis caprifoliacearum, and to reduced P. khasiana to synonymy. Braun (1992) examined type material of Pseudocercospora khasiana and compared it with P. viburni-cylindrici, described from China. The two species are morphologically very similar. Therefore, Braun (1992) proposed to reduce P. khasiana to synonymy with P. viburni-cylindrici, a treatment followed by Crous & Braun (2003) and Kamal (2010). Differences between Chinese and Indian collections were considered to be variation within a single species. The Chinese type material of P. viburni-cylindrici is characterised by having loose to often dense or even coremioid conidiophore fascicles, similar to P. caprifoliacearum, but narrower conidia, 2–4.5 µm wide. We prefer to maintain P. caprifoliacearum (inclusing P. khasiana) as separate species, at least for the interim until cultures and molecular data will be available.

Pseudocercospora depazeoides (Desm.) U. Braun & Crous, comb. nov.

MycoBank MB814570 (Fig. 49)

Fig. 49
figure49

Pseudocercospora depazeoides (Rabenh., Herb. Viv. Mycol., no. 587, lectotype of Passalora penicillata, HAL). A. Conidiophore fascicles. B. Conidia. Bar = 10 µm.

Basionym: Exosporium depazeoides Desm., Ann. Sci. Nat., Bot., sér. 3, 11: 364 (1849).

Synonyms: Passalora penicillata Ces., in Rabenh., Herb. Viv. Mycol., no. 587 (1857), with description on label [lectotype (designated here, MycoBank, MBT202789): Italy: Vercellis, on Sambucus nigra, 1856, V. de Cesati, Rabenh., Herb. Viv. Mycol. 587 (HAL)].

Cercospora penicillata (Ces.) Fresen., Beitr. Mykol.: 93 (1863).

Cercospora depazeoides (Desm.) Sacc., Mycoth. Ven., Cent. III, no. 280 (1875).

Cercospora sambucina Ellis & Kellerm., Amer. Naturalist 17: 1166 (1883) [lecotype (designated here, MycoBank, MBT202790): USA: Ohio, on Sambucus canadensis, Jul. 1883, W A. Kellerman 401 (NY 2343036); isolectotypes: CUP 41160, NY 2343037].

Cercospora depazeoides var. sambucina (Ellis & Kellerm.) Sacc., Syll. Fung. 4: 469 (1886).

Cercospora ticinensis Cavara, in Briosi & Cavara, Funghi Paras. Piante Colt. Util. Ess., no. 336 (1900) [lectotype (designated here, MycoBank, MBT202792): Italy: Pavia, Botanic Garden, on Sambucus nigra, Briosi & Cavara, Funghi Paras. Piante Colt. Util. Ess. 336 (HAL); isolectotypes: Briosi & Cavara, Funghi Paras. Piante Colt. Util. Ess. 336, e.g. BPI 441927, CUP, FH, K, MICH 15376, OSU 35059, PAD].

Cercospora depazeoides var. amphigena Sousa da Câmara, Revista Agron. 1: 59 (1903) [type: Portugal: Chaves (Traz-os-Montes), on Sambucus nigra, Dec. 1902, A. Pereira].

Cercospora depazeoides var. gagrensis Elenkin & Ohl, Bolez. Rast. 6: 108 (1912) [type: Georgia: Abkhazia: Gagra [Gagry], upper waterfall Zhoekvary River, on Sambucus nigra, 9 Aug. 1912, A. Elenkin (LE 158291)].

Cercospora sambuci F. Stevens & King, Illinois Biol. Monogr. 11: 59 (1927) [holotype: Costa Rica: Cartago, on Sambucus canadensis [mexicana], 7 Jul. 1923, F. L. Stevens (ILL 15175); isotypes: CUP 14660, NY 2343299].

Phaeoramularia penicillata (Ces.) X.J. Liu & Y.L. Guo, Acta Phytopathol. Sin. 12: 13 (1982).

Pseudocercospora sambucigena U. Braun, Crous & K. Schub., Mycotaxon 92: 400 (2005) [holotype: USA: Pennsylvania: Dauphon County, on Sambucus pubens, 21 Aug. 1921, O. E. Jennings, Acc. 6736 (NY 142641)].

Literature: Saccardo (1886: 468–469), Lindau (1910: 134), Gonzáles Fragoso (1927: 252), Vassiljevsky & Karakulin (1937: 235), Chupp (1954, 100, 103), Katsuki (1965: 16), Ellis (1976: 247), Braun & Mel’nik (1997: 55), Crous & Braun (2003: 156), Guo et al. (2005: 279–280), Bakalova & Borisova (2010: 39–40), Crous et al. (2013: 105–106).

Illustrations: Briosi & Cavara, Funghi Paras. Piante Colt. Util. Ess. 336, figs 13, Ellis (1976: 246, fig. 185 D), Braun & Crous (2005: 402, fig. 6), Bakalova & Borisova (2010: 136, pl. 4), Crous et al. (2013: 106, fig. 60).

Exsiccatae: Briosi & Cav., Funghi Paras. Piante Colt. Util. Ess. 336. Cavara, Fungi Longobard. Exs. 245, 984. Ellis & Everh., Fungi Columb. 599. Ellis & Everh., North Amer. Fungi 1749. Kabat & Bubak, Fungi Imperf. Exs. 97. Kerner von Maurilaun, Fl. Exs. Austr.-Hung. 788. Krieger, Fungi Saxon. Exs. 898, 899. Krieger, Schädl. Pilze Kulturgew. 193, 898. Krypt. Exs. 727. Linh., Fungi Hung. Exs. 294. Petr., Fungi Alban. Bosn. Exs. 7. Petr., Fl. Bohem. Morav. Exs. Pilze 508. Petr., Fungi Polon. Exs. 19. Petr., Mycoth. Gen. 1414. Poelt & Scheuer, Reliqu. Petr. 943. Rabenh., Fungi Eur. Exs. 879, 2070. Roum., Fungi Sel. Gall. Exs. 158, 3596. Sacc., Mycoth. Ven. 280. D. Sacc., Mycoth. Ital. 199, 1587. Săvul., Herb. Mycol. Rom. 195. Scheuer, Mycoth. Graec. 433. Seym. & Earle, Econ. Fungi 476. Syd., Mycoth. March. 1290, 2177. Syd., Mycoth. Germ. 1045, 1774, Thüm., Mycoth. Univ. 984. Vill, Fungi Bav. 996. Zahlbruckner, Krypt. Exs. 727 A, B.

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–10 mm diam, sometimes confluent and larger, at first pale greenish or greyish green, later brownish, finally grey to greyish white, often zonate, margin narrow, often formed as marginal line, somewhat raised, dark olivaceous-brown to blackish or dark purplish violet. Caespituli amphigenous, punctiform, dark brown to blackish, scattered. Mycelium internal; hyphae branched, septate, 2–4 µm wide, subhyaline to pale olivaceous, thin-walled. Stromata well-developed, substomatal to intraepidermal, subglobose to somewhat irregular, 10–100 µm diam, immersed to somewhat erumpent, medium to dark brown, composed of swollen hyphal cells, 3–7 µm diam. Conidiophores in small to large fascicles, loose to very dense, occasionally even almost coremioid, arising from stromata, emerging through stomata or erumpent, erect, straight, subcylindrical to flexuous, somewhat geniculate-sinuous, unbranched, 20–200(− 300) × 3–7 µm, short conidiophores 0–1-septate, longer ones pluriseptate throughout, pale olivaceous to dark olivaceous-brown or brown, tips often paler, thin-walled, smooth or almost so; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, 10–30 µm long, sympodially or occasionally percurrently proliferating, conidiogenous loci inconspicuous, unthickened or almost so and not darkened to subconspicuous by being subdenticulate, tips truncate or subtruncate, 1.5–2.5 µm wide, or slightly refractive to darkened-refractive, in front view occasionally paracercosporoid, visible as minute circle, 1.5–2 µm diam. Conidia solitary, obclavate-cylindrical, straight to curved, 30–140(− 155) × (3.5−)4–6.5(− 7.5) µm, (1−)2–8(− 9)-septate, hyaline, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, occasionally somewhat rough-walled, apex obtuse, base short to occasionally long obconically truncate, (1.5−)2–2.5(− 3) µm wide, hila unthickened or almost so, not darkened to somewhat refractive or slightly darkened-refractive.

Culture characteristics: Colonies on MEA reaching 16 mm diam after 30 d in the dark at 24°C, circular to subcircular, with smooth to slightly irregular margin, prominently convex, with moderate aerial mycelium, pale greenish grey to pale olivaceous-grey (surface) and olivaceous-black (reverse).

Lectotype (designated here, MycoBank, MBT202793): France: on Sambucus nigra, autumn, J. B. H. J. Desmazières, Desm., Pl. Crypt. France 1849 (NY 2343300). Isolectotypes: Desm., Pl. Crypt. France 1849, e.g. FH, G, PC. Epitype (designated here, MycoBank, MBT202794). The Netherlands: Milingerwaard, on Sambucus nigra, 2007, P. W. Crous (CBS H-20391). Ex-epitype culture: CBS 126000.

Host range and distribution: On Sambucus (canadensis [intermedia var. insularis, mexicana], chinensis, ebulus, glauca, kamtschatica, nigra, pubens, racemosa, williamsii [buergeriana], Sambucus sp.), Adoxaceae, Asia (China, Japan, Russia, Far East), Australia, Caucasus (Armenia, Azerbaijan, Georgia), Europe (Austria, Belgium, Bulgaria, Cyprus, Czech Republic, Denmark, Germany, Greece, Hungary, Italy, Latvia, Netherlands, Poland, Portugal, Romania, Russia, Slovakia, Spain, UK, Ukraine, former Yugoslavia), New Zealand, Central America and West Indies (Costa Rica, Haiti), North America (Canada; Mexico; USA, Connecticut, Florida, Illinois, Iowa, Kansas, Louisiana, Maine, Maryland, Mississippi, New York, New Jersey, Ohio, Pennsylvania, South Carolina, Washington, West Virginia, Wisconsin).

Notes: The generic affinity of Cercospora depazeoides has been confused and misinterpreted. Owing to the structure of the conidiogenous loci, ranging from inconspicuous to more conspicuous by being very slightly thickened or somewhat darker than the surrounding wall of the conidiogenous cell by being refractive, sometimes also visible as minute circle in front view (paracercosporoid), as well as usually colourless or almost so to pale olivaceous conidia, Crous & Braun (2003) maintained C. depazeoides as species of Cercospora s. str., and Braun & Crous (2005) introduced Pseudocercospora sambucigena for collections with quite indistinct conidiogenous loci. The variability and morphological range of the conidiogenous loci of this species are reminiscent of the locus characters of Phaeoisariopsis griseola (now Pseudocercospora griseola) and the species of the Paracercospora complex (Braun et al. 2013, Crous et al. 2013). Re-examinations of types and numerous other collections involved led to a reassessment of cercosporoid fungi on Sambucus spp. It emerged very clearly that a single species is involved which belongs in Pseudocercospora in the current sense and circumscription of this genus (Braun et al. 2013, Crous et al. 2013). Pseudocercospora sambucigena must be reduced to synonymy with P. depazeoides. The phylogenetic position of this species in the Pseudocercospora clade has recently been confirmed by Crous et al. (2013).

Lectotype material of Exosporium depazeoides contains besides the typical cercosporoid hyphomycete also an alternarioid fungus. It cannot be excluded that the cercosporoid as well as the alternarioid hyphomycete were included in the very meagre original description. However, we follow Saccardo’s treatment on the occasion of his introduction of the combination Cercospora depazeoides and confine this name to the cercosporoid element. Saccardo’s combination is usually cited as “Nuovo Giorn. Bot. Ital. 8: 187 (1876)” but it was first validly published in Mycotheca Veneta 280 (1875). The designated epitype, the ex-epitype culture and sequences obtained from this material help to fix the phylogenetic position and affinity of this species. The reallocation of C. depazeoides to Pseudocercospora has serious consequences on genetic level since Cercospora penicillata (i.e. C. depazeoides) being the type species of the genus Cercospora (Braun 1995, Braun et al. 2013). Thus, the allocation of C. depazeoides to Pseudocercospora, based on morphological reassesments and results of molecular sequence analyses, renders Cercospora an older heterotypic synonym of Pseudocercospora, which is undesirable and requires a proposal to conserve Cercospora with C. apii as conserved type under Art. 14.9, which is being made.

Eriksson, Fungi Paras. Scand. Exs. 42, deposited at HAL, has been examined and turned out to be a mixture of Ramularia sambucina and Cladosporium herbarum.

Pseudocercospora ebulicola (W. Yamam.) Deighton, Mycol. Pap. 140: 143 (1976). (Fig. 50)

Fig. 50
figure50

Pseudocercospora ebulicola (CUP-039732, lectotype). A. Conidiophore fascicle. B. Solitary conidiophore arising from a superficial hypha. C. Conidiophore tips. D. Conidia. Bar = 10 µm.

Basionym: Cercospora ebulicola W. Yamam., Trans. Sapporo Nat. Hist. Soc. 13: 139 (1934).

Literature: Chupp (1954: 100), Hsieh & Goh (1990: 50), Guo & Hsieh (1995: 47–48), Guo et al. (1998: 60–61), Crous & Braun (2003: 168).

Illustrations: Hsieh & Goh (1990: 50, fig. 34), Guo & Hsieh (1995: 49, fig. 46), Guo et al. (1998: 60, fig. 46).

Description: Leaf spots amphigenous, indefinite discolorations, yellowish to brownish, or circular to irregular spots, 1–8 mm diam, pale brown, yellowish brown to brown, later greyish white, margin indefinite or brown on the upper surface. Caespituli hypophyllous, effuse, olivaceous to brownish, forming irregular patches. Mycelium internal or partly external; superficial hyphae branched, 2–3 µm wide, septate, pale olivaceous. Stromata lacking or very small, only a few substomatal swollen hyphal cells, brown. Conidiophores in loose fascicles, 3–15, arising from internal hyphae or small hyphal aggregations, through stomata, erect, subcylindrical, curved, geniculate-sinuous to somewhat tortuous, simple or branched, 50–190 × 4–5.5 µm, 2–8-septate, olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci inconspicuous or subdenticulate, but always unthickened, not darkened. Conidia solitary, obclavate-cylindrical, straight to curved, 20–130 × 3–5 µm, 2–11-septate, subhyaline, yellowish to pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base subtruncate to short obconically truncate, 1.5–2.5 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202795): Taiwan: Sozan, on Sambucus javanica, 3 Dec. 1933, W. Yamamoto (CUP-039732). Isolectotype: HMAS 5197, K(M) IMI 7791. Topotypes: 20 Nov. 1933 (BPI 436018, 436019).

Host range and distribution: On Sambucus (javanica [formosana]), Adoxaceae, Asia (China, Sichuan, Zhejiang; Japan, Taiwan).

Notes: Chupp (1954) reduced Cercospora ebulicola to synonym with C. catenospora which is, however, incorrect. Cercospora ebulicola is a species of the genus Pseudocercospora whereas C. catenospora is a Phaeoramularia-like species of Passalora.

Pseudocercospora opuli (Fuckel) U. Braun & Crous, Mycophaerella and Anam.: 299 (2003). (Fig. 51)

Fig. 51
figure51

Pseudocercospora opuli (Fuckel, Fungi Rhen. Exs. 118, HAL, lectotype). A. Conidiophore fascicles and solitary conidiophore. B. Conidiophore tips. C. Conidia. Bar = 10 µm. U

Basionym: Cercospora penicillata f. opuli Fuckel, Fungi Rhen. Exs. 118 (1863).

Synonyms: Cercospora opuli (Fuckel) Höhn., in Kabát & Bubák, Fungi. Imperf. Exs. 445 (1907).

Cercospora viburni Sacc., in Sydow, Mycoth. March. 2773 (1889), nom. nud. (ICN 38.1).

Pseudocercospora viburni U. Braun, Nova Hedwigia 55: 219 (1992) [holotype: China: Hubei: Shennongjian, on Viburnum opulus subsp. calvescens [sargentii var. calvescens], 5 Aug. 1984, Y. L. Guo 266 (HMAS 47828)].

Pseudophaeoramularia opuli (Fuckel) U. Braun, Trudy Bot. Inst. im. V.L. Komarova 20: 19 (1997).

Passalora viburni-sargentii Y.L. Guo, Mycosystema 31: 161 (2012) [holotype: China: Hubei: Shennongjian, on Viburnum opulus subsp. calvescens [sargentii var. calvescens], 5 Aug. 1984, Y. L. Guo 266 (HMAS 47828)].

Misapplied names: Cercospora penicillata sensu Saccardo (1886: 468) and Chupp (1954: 103).

Phaeoramularia penicillata (Ces.) X.J. Liu & Y.L. Guo, Acta Phytopathol. Sin. 12(4): 13 (1982).

Literature: Saccardo (1886: 468), Lindau (1910: 136), Ferraris (1910: 136), Vassiljevsky & Karakulin (1937: 236), Chupp (1954: 103), Ellis (1976: 247), Braun & Mel”nik (1997: 19, 77).

Illustrations: Chupp (1954: 103), Ellis (1976: 248, fig. 186A), Braun (1992: 216, fig. 9), Braun & Mel”nik (1997: appendix, fig. 82), Guo (2012: 161, fig. 2).

Exsiccatae: Fuckel, Fungi Rhen. Exs. 118. Kabát & Bubák, Fungi Imperf. Exs. 445. Liro, Mycoth. Fenn. 295. Petr., Mycoth. Gen. 1312. Săvul., Herb. Mycol. Rom. 432. Scheuer, Mycoth. Graec. 382. Smarods, Fungi Lat. Exs. 448.

Description: Leaf spots amphigenous, subcircular to somewhat angular-irregular, 1–8 mm diam, sometimes confluent and larger, occasionally somewhat zonate, dull greenish, ochraceous to brown, centre later greyish to greyish white, with narrow dark margin. Caespituli amphigenous, punctiform, scattered, dark brown to blackish. Mycelium internal. Stromata substomatal or immersed, 10–80 µm diam, brown, composed of swollen hyphal cells, 2–7 µm diam, walls somewhat thickened. Conidiophores in small to fairly large fascicles, loose to dense, arising from stromata, emerging through stomata or erumpent, erect, straight, subcylindrical to moderately geniculate-sinuous, unbranched or occasionally with a single branchlet, 15–70(− 80) × (2.5−)3–7(− 8) µm, aseptate or sparingly septate, usually with 0–3 septa, at first subhyaline and thin-walled, later pale olivaceous to olivaceous-brown and walls somewhat thickened, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, about 10–40 µm long; conidiogenous loci inconspicuous to subconspicuous by being truncate and somewhat refractive or darkened-refractive, but unthickened, in front view sometimes visible as minute circles, about 1.5–2.5 µm diam. Conidia solitary to catenate, in simple, short acropetal chains or disarticulating, obclavate-subcylindrical to cylindrical, straight to somewhat curved, (20−)30–90(− 145) × (2.5−)3–6(− 7) µm, 1–10-septate, occasionally somewhat constricted at the septa, hyaline, subhyaline, later pale olivaceous, thin-walled, smooth, apex of primary and solitary conidia obtuse, rounded, truncate to short conically truncate in catenate conidia, base truncate to short obconically truncate, 2–3 µm wide, hila unthickened, not to slightly refractive or darkened-refractive.

Lectotype (designated by Braun & Mel”nik 1997: 20): Germany: Rheinland-Pfalz: “in sylva Hostrichiensi”, on Viburnum opulus, summer 1863 [Fuckel, Fungi Rhen. Exs. 118] (HAL). Isolectotypes: Fuckel, Fungi Rhen. Exs. 118, (FH, G, K(M), etc).

Host range and distribution: On Viburnum (edule [pauciflorum], opulus subsp. opulus, opulus subsp. calvescens [opulus var. calvescens, sargentii var. calvescens], opulus subsp. trilobum [opulus var. americanum, trilobum]), Adoxaceae, Asia (China, Kazakhstan, Russia, Turkey), Caucasus (Armenia, Azerbaijan, Georgia), Europe (Austria, Belarus, Bulgaria, Czech Republic, Denmark, Estonia, Finland, Germany, Great Britain, Italy, Latvia, Poland, Portugal, Romania, Russia, Slovakia, Sweden, Ukraine), Central America (Costa Rica), North America (Canada; USA, Idaho, Iowa, Kansas, Mississippi, Oklahoma, Wisconsin).

Notes: A record of this species on V. burejaeticum from the Far East of Russia (Egorova 2007) is uncertain and unproven and might rather pertain to P. varia. Viburnum burejaeticum does not belong to Viburnum sect. Opulus, but P. opuli seems to be confined to species of this section, which has been confirmed as well supported clade in phylogenetic studies (Winkworth & Donoghue 2005, Clement et al. 2014). Viburnum lantana and V. orientale (Crous & Braun 2003) are additional unproven and doubtful hosts of P. opuli not belonging to sect. Opuli. The taxonomic history of this species is complicated and characterised by confusions and misinterpretations. Passalora penicillata (syn. Cercospora penicillata, now Pseudocercospora depazeoides) was introduced for a cercosporoid fungus on Sambucus nigra. Fuckel (1863) added C. penicillata f. opuli, and Fuckel (1870) treated the fungus on Viburnum opulus as sole species of C. penicillata, i.e. “f. penicillata” on Sambucus nigra was not mentioned, which seems to be the reason for Saccardo’s (1886) misinterpretation of the name C. penicillata. Chupp (1954) followed Saccardo’s misinterpretation, although the confusion of the names involved had already been discussed and corrected by Lindau (1910). The proposed combination Phaeoramularia penicillata (Liu & Guo 1982) was also based on Chupp’s wrong interpretation of the name Cercospora penicillata. Höhnel’s name C. opuli can be interpreted as new species (see Braun & Mel’nik 1997, Crous & Braun 2003) or as new combination at new rank according to Art. 41.4. We prefer to follow the second version. The morphology of this species is unusual and caused additional confusion. The conidia are formed singly as well as in chains or disarticulate in smaller units. The conidiogeous loci range from being inconspicuous to subconspicuous by being somewhat refractive or darkened-refractive, but they are consistently unthickened. Braun (in Braun & Mel’nik 1997) classified loci and hila of this species as intermediate between Passalora and Pseudocercospora and proposed the new genus Pseudophaeoramularia for such species. However, based on results of molecular sequence analyses and morphological reassessments, Pseudophaeoramularia was reduced to synonymy with Pseudocercospora and C. opuli was reallocated to the latter genus (Crous & Braun 2003, Braun et al. 2013, Crous et al. 2013).

Pseudocercospora viburni and Passalora viburni-sargentii are homotypic synonyms. Chinese material on Viburnum opulus subsp. calvescens has been examined and found to be indistinguishable from European material of P. opuli.

A collection of this species from the Asian part of Turkey has been examined (Kütahya, on Viburnum opulus, 9 Jul. 1953, ex herb. Petrak, GZU).

Pseudocercospora varia is morphologically very close to and confusable with P. opuli, but the conidia are formed singly. The latter species seems to be confined to species on Viburnum sect. Opuli. The taxonomic meaning of conidial catenation, which occurs in varying degrees in P. opuli, is not quite clear and has to be elucidated by means of molecular methods. Other morphological characters of P. opuli and P. varia are very similar. The two species are treated as separate species, at least tentatively.

Pseudocercospora tinea Y.L. Guo & W.H. Hsieh, Mycosystema 7: 124 “1994” (1995). (Fig. 52)

Fig. 52
figure52

Pseudocercospora tinea (based on Guo 1995: 125, fig. 4). A. Conidiophore fascicles. B. Conidiophores. C. Solitary conidiophores arising from superficial hyphae. D. Conidia. Bar = 10 µm.

Misapplied name: Stigmina tinea sensu Hsieh & Goh 1990.

Literature: Hsieh & Goh (1990: 51–52, as Stigmina tinea), Guo et al. (1998: 63–64).

Illustrations: Hsieh & Goh (1990: 53, fig. 36, as Stigmina tinea), Guo (1995: 125, fig. 4), Guo et al. (1998: 64, fig. 49).

Description: Leaf spots amphigenous, subcircular to angular-irregular and vein-limited, 0.5–7 mm diam, often confluent and along the leaf margin, brown to dark brown, margin indefinite or centre later greyish brown, dingy grey to greyish white, margin dark brown above and paler brown below, sometimes with diffuse yellowish to greyish brown halo. Caespituli amphigenous, punctiform, dark brown. Mycelium internal and external; superficial hyphae hypophyllous, emerging through stomata or arising from the base of conidiophore fascicles, branched, septate, pale olivaceous, thin-walled, smooth, 1.5–2.5 µm wide. Stromata well-developed, substomatal, 15–75 µm diam, brown. Conidiophores in small to moderately large fascicles, divergent to usually dense, arising from stromata, through stomata or solitary, arising from superficial hyphae, erect, straight to curved, subcylindrical, somewhat attenuated towards the tip or irregular in width, geniculate-sinuous, unbranched, relatively short, 5–35 × 2–5 µm, 0–1-septate, pale olivaceous, olivaceous-brown to brownish, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores often reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous or subdenticulate, but always unthickened and not darkened. Conidia solitary, obclavate-cylindrical, 10–120 × 2–3.5(− 4) µm, 1–11-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base subtruncate to usually obconically truncate, 1–2.5 µm wide, hila unthickened, not darkened.

Holotype: China: Guangdong Province: Guangzhou, on Viburnum macrocephalum, 6 Nov. 1961, Q. M. Ma & X. J. Liu 1079 (HMAS 67254).

Host range and distribution: On Viburnum (luzonicum, macrocephalum, plicatum var. tomentosum [tomentosum], suspensum, Viburnum sp.), Adoxaceae, Asia (China, Anhui, Guangdong, Sichuan, Zhejiang; Japan, Taiwan).

Notes: Guo and Hsieh (in Guo 1995) compared this species with Cercospora tinea, supposed that the two species could be synonymous, but hesitated to reduce them to synonymy. Therefore, they did not propose a new combination based on Cercospora tinea, but preferred to describe a new species with a Chinese type collection. Hence, the name Pseudocercospora tinea is valid. However, as already stated in Braun & Hill (2002), Cercospora tinea (now Pseudocercospora viburnigena) is clearly distinct from the Chinese species by having non-geniculate, only percurrently proliferating conidiogenous cells with fine annellations. Superficial hyphae are only rarely formed. Pseudocercospora tinea is possibly rather common in China and Taiwan. Two additional Taiwanese collections on Viburnum sp. have been examined and identified by C. Nakashima (TUA 40, 56), and Japanese collections on Viburnum suspensum and V. plicatum var. tomentosum [V. tomentosum] have been revised and confirmed as P. tinea by C. Nakashima (CNS567, CNS976).

Pseudocercospora varia (Peck) J.K. Bai & M.Y. Cheng, Acta Mycol. Sin. 11: 123 (1992). var. varia (Fig. 53a)

Fig. 53a
figure53a

Pseudocercospora varia var. varia (BPI 442161, isotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora varia Peck, Rep. (Annual) New York State Mus. Nat. Hist. 35: 141 (1884).

Literature: Saccardo (1886: 468), Chupp (1954: 105), Guo & Hsieh (1995: 51), Crous & Braun (2003: 418).

Exsiccatae: Clements & Clements, Crypt. Format. Colorad. 280. Ellis & Everh., N. Amer. Fungi 3190.

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–8 mm diam, sometimes confluent and larger, pale to medium dark brown or reddish brown, later grey to greyish white, margin indistinct, formed as marginal line or with narrow to moderaterly wide, dark brown margin, sometimes somewhat raised, and sometimes with ochraceous, yellowish brown to brownish halo. Caespituli amphigenous, punctiform, scattered, brown, dark brown to almost blackish. Mycelium internal (in some collections with superficial hyphae probably formed by conidial germination); hyphae branched, septate, pale, 1.5–3 µm wide, thin-walled, smooth. Stromata substomatal or intraepidermal, small to well-developed, 10–60 µm diam, subglobose, pale to medium brown or yellowish brown, cells globose or subglobose to somewhat irregular, 3–10 µm diam, wall thickened, to 1 µm. Conidiophores solitary or in usualy small fasciles, mostly 2–15, usually divergent, rarely dense, arising from stromata, through stomata or erumpent, erect, straight to somewhat curved, subcylindrical or attenuated towards the tip, mostly not geniculate-sinuous, occasionally somewhat geniculate-sinuous in the upper half, unbranched, (5−)10–80(− 100) × 2–9 µm, usually 0–3-septate, at first very pale, hyaline to subhyaline, later pale olivaceous, olivaceous-brown or yellowish brown, above all below, mostly paler towards the tip, wall thin to usually somewhat thickened, to 0.5 µm, above all below, sometimes to 1 µm thick, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 15–40 µm long, conidiogenous loci inconspicuous or visible by being truncate or subdenticulate, but always unthickened and not darkened, occasionally somewhat refractive or loci visible as minute circle. Conidia solitary, obclavate-cylindrical, cylindrical, rarely subacicular, straight to somewhat curved, (15−)20–85(− 90) × 2–6.5 µm, 1–7-septate, hyaline to subhyaline, thin-walled, smooth, apex obtuse, occasionally subacute, base truncate to short or long obconically truncate, 1–2.5 µm wide, hila unthickened, not darkened.

Holotype: USA: New York: Albany County, East Berne, on Viburnum acerifolium, Aug., C. H. Peck (NYS). Isotype: BPI 442161.

Host range and distribution: On Viburnum (acerifolium, alnifolium, cassinoides, dilatatum, lentago, odoratissimum, prunifolium, pubescens, rafinesquianum var. affine [affine], suspensum, Adoxaceae, North America (Canada; USA, Alabama, Alaska, Florida, Idaho, Illinois, Iowa, Kansas, Massachusetts, Mississippi, New York, Oklahoma, Oregon, South Carolina, Texas, Vermont, Virginia, West Virginia, Wisconsin, Wyoming).

Notes: Collections on Viburnum edule [V. pauciflorum], V. opulus and V. trilobum have a tendency to form conidia in short chains and short lateral branchlets. They are referred to as P. opuli. Records of this species from New Zealand (Gagdil 2005) on Viburnum lantata and V. carlesii are quite unclear. The collections concerned have been examined (PDD 19826, 19843, 1985), but sufficient fructification for a final identification has not been found. Superficial hyphae with solitary conidiophores found on “V. lantana” are not in accordance with P. varia. The host identity is also uncertain (short petioles do not coincide with V. lantana but they are rather in favour of V. burejaeticum. Other records of C. varia on Lonicera spp. are also based on misidentifications. Collections on L. japonica belong to Pseudocercospora lonicericola (material examined: BPI 442155), other specimens, e.g. on L. ciliata (BPI 442168–442170), refer to Passalora antipus (Ellis & Holw.) U. Braun & Crous.

var. viburni-sargentii U. Braun & H.D. Shin, var. nov. MycoBank MB814558 (Fig. 53b)

Fig. 53b
figure53b

Pseudocercospora varia var. viburni-sargentii (KUS 14011, holotype). A. Conidiophore fascicles. B. Conidia. Bar = 10 µm.

Literature: Guo & Hsieh (1995: 51, as Pseudocercospora varia), Guo et al. (1998: 65, as Pseudocercospora varia), Kim & Shin (1999), Shin & Kim (2001: 226, as Pseudocercospora varia).

Illustrations: Guo & Hsieh (1995: 51, fig. 48), Guo et al. (1998: 66: 50), Shin & Kim (2001: 227, fig. 104).

Diagnosis: Morphologically close to North American collections of var. varia, but characterised by having conidiophores in larger, dense fascicles, about 10–30; conidia, 35–145 × 3.5–6.5 µm, 2–15-septate.

Holotype: Korea: Pyeongchang, on Viburnum sargentii, 25 Aug. 1997, H. D. Shin (KUS-F 14011).

Host range and distribution: On Viburnum sargentii, Adoxaceae, Asia (China, Heilongjian, Liaoning; Korea).

Notes: The Asian collections on Viburnum sargentii from China and Korea differ from North American specimens in having much larger, dense fascicles of conidiophores. The taxonomic status of the Asian collections is not quite clear and in urgent need of molecular confirmation. It cannot be excluded that this is a separate species, but for the interim we prefer a conservative treatment as variety.

Pseudocercospora viburnicola U. Braun, sp. nov. MycoBank MB814559 (Fig. 54)

Fig. 54
figure54

Pseudocercospora viburnicola (BPI 442174, holotype). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Diagnosis: Differs from Pseudocercospora viburni-nudi in having much longer, usually obclavate conidia, 25–125 × 4.5–8 µm, with 2–10 septa and somewhat thickened walls and subconspicuous conidiogenous loci. Pseudocercospora caprifoliacearum, known from India on Viburnum sp., has longer conidiophores, 60–155 µm, and the conidia are 3–6.5 µm wide and thin-walled.

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–8 mm diam, medium to dark brown, finally with pale centre, greyish brown to dingy greyish white, margin indefinite or narrow, darker, or surrounded by darker veins, occasionaly somewhat raised. Caespituli amphigenous, punctiform, scattered, blackish. Mycelium internal. Stromata lacking or substomatal to intraepidermal, 10–50 µm diam, brown, swollen hyphal cells 2–7 µm diam, walls slightly thickened. Conidiophores in small, divergent fascicles, mostly 3–12, arising from substomatal hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to distinctly geniculate or geniculate-sinuous in the upper fertile portion, unbranched, 20–100 × 3.5–8(− 10) µm, (0−) 1–4(− 5)-septate, medium to dark brown throughout or somewhat paler towards the tip, wall somewhat thickened, to 0.8 µm, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, proliferation sympodial, occasionally with enteroblastic, monopodial proliferation leaving coarse annellations, conidiogenous loci inconspicuous to conspicuous by being denticle-like, 1.5–3 µm diam, often somewhat refractive or even appearing darkened, in front view sometimes visible as minute circle with dark rim (caused by the relatively thick and dark walls of the conidiophores and conidiogenous cells), but loci not thickened. Conidia solitary, obclavate, often with almost rostrate apex, shorter conidia subcylindrical, 25–125 × 4.5–8 µm, 2–10-septate, occasionally subhyaline to pale olivaceous, but usually medium olivaceous or olivaceous-brown, wall thin to somewhat thickened (to 0.8 µm), smooth, apex obtuse to subacute, base obconically truncate, 2–3 µm wide, hila unthickened, not darkened, but often somewhat refractive or ultimate rim slightly darker.

Holotype: USA: Wisconsin: Madison, Arboretum, on Viburnum sp. [? rafinesquianum] (as Viburnumlentago”), 5 Sep. 1953, W W. Diehl (BPI 442174).

Host range and distribution: On Viburnum (cassinoides, Viburnum sp.), Adoxaceae, North America (USA, Wisconsin, West Virginia).

Notes: An additional collection of this species has been examined [USA: West Virginia: Pocahontas County, on Viburnum cassinoides, 6 Aug. 1909, B. Brooks, hb. J. L. Sheldon 3815 (BPI 439059)]. This species is easily distinguishable from all Pseudocercospora species on hosts of the Adoxaceae by its consistently solitary, pigmented, much wider conidia with somewhat thickened walls. Other comparable cercosporoid fungi on Viburnum spp., e.g. Pseudocercospora varia, are quite distinct by narrower, mostly subhyaline conidia and unthickened conidiogenous loci and conidial hila. The shape of the conidia with somewhat thickened walls and the structure of the conidiogenous loci in this species are reminiscent of former Prathigada species which have turned out to belong to Pseudocercospora (Braun et al. 2013).

Pseudocercospora viburni-cylindrici (F.L. Tai) U. Braun, Nova Hedwigia 55: 219 (1992). (Fig. 55)

Fig. 55
figure55

Pseudocercospora viburni-cylindrici (HMAS 01929, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercospora viburni-cylindrici F.L. Tai, Syll. Fung. Sin.: 907 (1979).

Synonyms: Cercospora viburnicola F.L. Tai, Lloydia 11: 54 (1948), nom. illeg. (Art. 53.1), non C. viburnicola W.W. Ray, 1941.

Pseudocercospora viburni-cylindrici (FL. Tai) Y.L. Guo & W.X. Zhao, Acta Mycol. Sin. 12: 198 (1993).

Literature: Guo & Hsieh (1995: 52), Guo et al. (1998: 65); Crous & Braun (2003: 421).

Illustrations: Tai (1948: 37, fig. 1), Braun (1992: 216, plate 2, fig. 10), Guo & Hsieh (1995: 53, fig. 49), Guo et al. (1998: 67, fig. 51).

Description: Leaf spots amphigenous, subcircular to angular-irregular, size variable, 1–8 mm diam, sometimes confluent, brown to dark brown or even blackish, margin indefinite or finally centre brown, greyish brown or greyish white with darker margin, brown, sometimes with a diffuse olivaceous halo. Caespituli hypophyllous, punctiform, scattered, effuse, dark, brown to dark brown. Mycelium internal. Stromata well-developed, subglobose to somewhat irregular, substomatal, 20–35 µm diam, brown. Conidiophores in small to moderately large, somewhat divergent to dense or even coremioid fascicles, arising from stromata, through stomata, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 50–150 × 3–4.5 µm, 3–7-septate, medium olivaceous-brown to brown, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, about 10–30 µm long, conidiogenous loci inconspicuous unthickened and not darkened. Conidia solitary, obclavate-cylindrical, straight or only slightly curved, 25–75 × 2.5–4 µm, 2–5-septate, pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: China: Yunnan Province: Kunming, on Viburnum cylindricum, Adoxaceae, Jun. 1938, Xu Ren (HMAS 01929).

Host range and distribution: Only known from the type collection.

Notes: Braun (1992) reduced Pseudocercospora khasiana to synonymy with P. viburni-cylindrici, although the conidia in the Indian type material are distinctly wider and the conidiophore fascicles are smaller and not coremioid. Therefore, we prefer to consider P. khasiana (= P. caprifoliacearum) a distinct species of its own, at least for the interim (see notes under the latter species). Korean material on Viburnum erosum, referred to as P. viburni-cylindrici by Shin & Kim (2001: 228), differs in having much longer, pluriseptate, cylindrical conidia with truncate base. The material concerned is described as a new species, Pseudocercospora viburni-erosi.

Pseudocercospora viburni-erosi U. Braun & H.D. Shin, sp. nov.

MycoBank MB814560 (Fig. 56)

Fig. 56
figure56

Pseudocercospora viburni-erosi (KUS-F14126, holotype). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Literature: Shin & Kim (2001: 228).

Illustration: Shin & Kim (2001: 229, fig 105).

Diagnosis: Differs from P. viburni-cylindrici in having large, dense, often coremoid conidiophore fascicles and above all much longer, pluriseptate, cylindrical-filiform conidia, length 50–130 µm, with truncate base.

Description: Leaf spots amphigenous, scattered to confluent, circular or almost so to angular-irregular, 3–10 mm diam, sometimes confluent and larger, to 15 mm diam, at first brown to dark brown, later with greyish brown to grey centre surrounded by a brown to dark brown border, finally centre turning greyish white, with raised brown border line. Caespituli hypophyllous, brown, punctiform. Mycelium internal; hyphae branched, septate, hyaline, 2–3 µm wide. Stromata substomatal, small to well-developed, 15–30 µm diam, globose or subglobose, brown. Conidiophores in well-developed fascicles, to 30, dense to very dense, coremioid, arising from stromata, through stomata, erect, straight, subcylindrical or only slightly geniculate-sinuous, above all in the upper portion, unbranched, 60–120 × 3–4 µm, 3–8-septate, olivaceous-brown to brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, cylindrical-Aliform, straight or almost so to moderately curved, 50–130 × 2.5–4 µm, 3–12-septate, subhyaline to very pale olivaceous, thin-walled, smooth, apex obtuse, base truncate or almost so, 2–2.5 µm wide, hila unthickened, not darkened.

Holotype: Korea: Kwachon, on Viburnum erosum var. taquetii, 4 Sep. 1997, H. D. Shin (KUS-F14126).

Host range and distribution: Only known from the type collection.

Notes: Type material of the new species was previously assigned to Pseudocercospora viburni-cylindrici (Shin & Kim 2001). The two species are morphologically very close, but Korean material on Viburnum erosum differs in having large, dense, often coremoid conidiophore fascicles and above all much longer, pluriseptate, cylindrical-Aliform conidia, length 50–130 µm, with truncate base. Based on these obvious differences, the Korean material on Viburnum erosum is better excluded from P. viburni-cylindrici and treated as separate species.

Pseudocercospora viburnigena U. Braun & Crous, Mycol. Progr. 1: 23 (2002). (Fig. 57)

Fig. 57
figure57

Pseudocercospora viburnigena (Sacc., Mycoth. Ven. 1252, HAL, lectotype). A. Conidiophore fascicles. B. Stroma cells. C. Conidiophores. D. Conidia. Bars = 10 µm.

Basionym: Cercospora tinea Sacc., Michelia 1: 268 (1878), non Pseudocercospora tinea Y.L. Guo & W.H. Hsieh 1994.

Synonyms: Cercoseptoria tinea (Sacc.) Deighton, Mycol. Pap. 140: 167 (1976).

Stigmina tinea (Sacc.) M.B. Ellis, More Dematiaceous Hyphomycetes: 118 (1976).

Cercostigmina tinea (Sacc.) U. Braun, Cryptog. Bot. 4: 108 (1993).

Literature: Chupp (1954: 105), Katsuki (1965: 16), Kirk (1999), Crous & Braun (2003: 421), Crous et al. (2013: 108).

Illustrations: Ellis (1976: 117, fig. 83B), Kirk (1999: fig., unnumbered), Crous et al. (2013: 109, fig. 65).

Description: Leaf spots amphigenous, subcircular to angular-irregular, sometimes vein-limited, 2–15 mm diam, pale to darker brown or centre later greyish brown to dingy grey, margin indefinite or darker, sometimes with reddish tinge, occasionally slighty raised. Caespituli amphigenous, punctiform, olivaceous to dark or blackish brown, scattered. Mycelium internal, external mycelium lacking or occasionaly with some external hyphae; hyphae branched, septate, 1.5–4 µm wide, subhyaline, thin-walled, smooth. Stromata substomatal or immersed, 15–80 µm diam, occasionally larger, to 120 µm diam (hypophyllous stromata smaller and substomatal, epiphyllous ones larger and immersed to somewhat erumpent), subglobose, brown to dark brown, composed of brown swollen hyphal cells, 3–6 µm diam, subglobose to somewhat irregular, walls thickened, 0.5–2 µm. Conidiophores in somewhat divergent to usually dense, sometimes very dense fascicles, larger fascicles sporodochial, arising from stromata, emerging through stomata or erumpent, occasionally with solitary conidiophores arising from superficial hyphae, straight to somewhat curved-sinuous, not geniculate, unbranched, cylindrical, subcylindrical or slightly attenuated towards the tip, sometimes ampulliform, apex at first rounded, later truncate or subtruncate, 5–30(− 50) × 2–5(− 6), 0(− 2)-septate, pale brown or olivaceous-brown, thin-walled, smooth, occasionally slightly rough-walled; conidiophores mostly reduced to conidiogenous cells, occasionally integrated, terminal, 5–20 µm long, with a single terminal conidiogenous locus, 2–3 µm wide, neither thickened nor darkened, proliferation percurrent, with 1–3 not very conspicuous annellations. Conidia solitary, acicular to narrowly cylindrical-filiform, occasionally somewhat obclavate, straight to curved or somewhat sigmoid, (20−)30–110(− 120) × 2–5 µm, (1−)3–11-septate, subhyaline to very pale olivaceous or somewhat brownish, thin-walled, smooth, guttulate when fresh, apex obtuse to subacute, base truncate to somewhat obconically truncate, 1.5–3 µm wide, hila unthickened, not darkened.

In vitro: Colonies on MEA reaching 23 mm diam after 30 d at 24°C in the dark, circular, convex, margin smooth, distinctly darker than the rest of of colony, slightly folding occurs towards the margin of the colony, with moderate to profuse aerial mycelium, surface olivaceous-grey, reverse greenish black.

Lectotype (designated here, MycoBank, MBT202797): Italy: Padova, on Viburnum tinus, Oct. 1877, Bizzozero [Sacc., Mycoth. Ven. 1252] (HAL, s.n.). Isolectotypes: Sacc., Mycoth. Ven. 1252, e.g. B, BPI 441941, FH, HBG. Epitype (designated in Crous et al. 2013: 108): The Netherlands: Bilthoven, Sweelincklaan 87, on Viburnum davidii, 26 May 2008, M. K. Crous (CBS H-20393). Ex-epitype culture: CBS 125998.

Host range and distribution: On Viburnum (davidii, plicatum var. plicatum, tinus, Viburnum sp.), Adoxaceae, Europe (Germany, Great Britain, Italy, The Netherlands, Portugal, Spain), North America (USA, Louisiana).

Notes: A lectotypification of Cercospora tinea is necessary. Type material or any other samples are not listed for PAD in Gola (1930). Therefore, one of the syntypes distributed as “Sacc., Mycoth. Ven. 1252” is designated as lectotype. The phylogenetic affinity of this species, previously referred to as Stigmina and Cercostigmina, was elucidated by Crous et al. (2013). Based on molecular sequence analyses, it could be demonstrated that this species clusters in a big Pseudocercospora clade.

Pseudocercospora tinea (see above) is quite distinct from Cercospora tinea by having superfical hyphae with solitary conidiophores and conidiogenous cells sympodially proliferating. Katsuki (1965) recorded Cercospora tinea from Japan on Viburnum dilatatum and V. suspensum. Kobayashi (2007) added V. davidii, V. erosum, V. plicatum and V. sieboldii as host species for Japan. Japanese collections on Viburnum suspensum and V. plicatum var. tomentosum [V. tomentosum] have been examined by C. Nakashima (CNS567, CNS976) and turned out to belong to P. tinea. Therefore, all other Japanese records of P. viburnigena [C. tinea] remain unproven and doubtful, i.e, they seem rather to refer to P. tinea.

A single North America collection has been examined and confirmed (USA: Louisiana: Lafayette, on Viburnum plicatum, 21 Sep. 1886, A. B. Langlois, BPI 441939, 441940).

Pseudocercospora viburni-nudi U. Braun, sp. nov. MycoBank MB814562 (Fig. 58)

Fig. 58
figure58

Pseudocercospora viburninudi (BPI 442176, holotype). A. Conidiophore fascicle. B. Conidiophore. C. Conidiophore tips. D. Conidia. Bar = 10 µm.

Diagnosis: Distinguished from all other species of Pseudocercosora on Viburnum spp. and other hosts of the Adoxaceae by having much shorter, 1–4(− 5)-septate conidia, 15–40 × 3–7 µm.

Description: Leaf spots amphigenous, subcircular to angular-irregular, sometimes vein-limited, 2–12 mm diam or confluent and larger, medium to dark brown, margin indefinite, later with paler centre, greyish brown, grey to greyish white, surroundced by a narrow to moderately wide darker border, dark brown to almost blackish, sometimes with diffuse reddish halo. Caespituli amphigenous, punctiform, scattered, dark brown to blackish. Mycelium internal. Stromata substomatal to intraepidermal, 15–45 µm diam, dark brown. Conidiophores in small to moderately large fascicles, 3–18, arising from stromata, through stomata or erumpent, divergent, erect, straight, subcylindrical, only terminal part somewhat geniculate-sinuous or subdenticulate, unbranched, 40–105 × 3–5 µm, base occasionally somewhat wider, to 7 µm, apex sometimes slightly swollen, to 8 µm wide, 1–5-septate, medium to dark brown throughout or paler towards the tip, wall somewhat thickened, to 1 µm, conidiophores occasionally with a single enteroblastic rejuvenation leaving a single conspicuous annellation; conidiogenous cells integrated, terminal, 15–40 µm long, sympodially proliferating, conidiogenous loci inconspicuous to conspicuous by being subdenticulate, denticle-like loci about 1.5–2 µm diam, but loci always unthickened and not darkened. Conidia solitary, short obclavate-cylindrical, fusiform, ellipsoid, straight to curved, 15–40 × 3–7 µm, 1–4(− 5)-septate, often slightly constricted at the septa, subhyaline to mostly pale to medium olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse, base short obconically truncate, sometimes rounded, 1.5–2.5(− 3) µm wide, hila unthickened, not darkened.

Holotype: USA: South Carolina: Murrells Inlet, on Viburnum nudum, Adoxaceae, 15 Oct. 1943, Carter 855 (BPI 442176).

Host range and distribution: Only known from the type collection.

Aizoaceae

Cercospora

Key to Cercospora species on Aizoaceae
  1. 1

    Conidia obclavate-cylindrical to acicular, to 125 µm long, base truncate to obconically truncate; on Trianthema portulacastrum ………. C. trianthematis

    Conidia acicular, to 200 µm long, base truncate; on Tetragonia tetragonoides ………. C. tetragoniae

Cercospora species on Aizoaceae

Cercospora tetragoniae (Speg.) Siemaszko, Mater. Mikol. Fitopatol. Rossii 1(3): 40 (1915). (Fig. 59)

Fig. 59
figure59

Cercospora tetragoniae (LPS 16153, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Cercosporina tetragoniae Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 20: 429 (1911).

Synonyms: Cercospora tetragoniae (Speg.) Vassiljevsky, in Vassiljevky & Karakulin, Fungi imperfecti parasitici 1. Hyphomycetes: 221 (1937).

Cercospora tetragoniae (Speg.) Chupp, in Viégas, Bol. Soc. Bras. Agron. 8: 54 (1945).

Literature: Chupp (1954: 27), Katsuki (1965: 7), Sutton & Pons (1980: 216), Braun & Mel’nik (1997: 97), Braun (2000b: 78), Crous & Braun (2003: 400).

Illustration: Sutton & Pons (1980: 207, fig. 1F).

Description: Leaf spots amphigenous, circular or subcircular, 2–10 mm diam, scattered to confluent and larger, up to 20 mm, pale brown, dingy olivaceous, greyish brown to grey or dingy greyish white, margin brown to reddish brown, occasionally somewhat zonate. Caespituli amphigenous, mostly epiphyllous, punctiform, dark brown to blackish. Mycelium internal. Stromata lacking or almost so to small, substomatal, 10–25(− 40) µm diam, brown. Conidiophores solitary or fasciculate, 2–10, divergent, arising from internal hyphae or stromata, through stomata or occasionally erumpent, erect, straight, subcylindrical to geniculate-sinuous, unbranched, 20–125 × 3–7 µm, 0–6-septate, pale yellowish, olivaceous-brown to brown, pale towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, sometimes conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 2.5–4 µm diam. Conidia solitary, acicular, straight to curved, 30–200 × 2.5–5 µm, 3- to pluriseptate, hyaline, thin-walled, smooth, apex pointed, base truncate, 2.5–4 µm wide, hila thickened and darkened.

Holotype: Argentina: La Plata, on Tetragonia tetragonoides, 18 Nov. 1909, C. Spegazzini (LPS 16153). Isotype: K(M) IMI 241730 (slide).

Host range and distribution: On Tetragonia (tetragonoides [expansa]), Aizoaceae, Africa (Cameroon, Kenya, Malawi, Sierra Leone, Tanzania, Uganda, Zimbabwe), Asia (Brunei, Israel, Japan), Caucasus (Georgia), Central and South America (Argentina, El Salvador, Brazil), North America (USA, Maryland, Indiana, Texas).

Notes: This species belongs to the C. apii s. lat. complex. Records from Brazil on Spinacia oleracea (Mendes et al. 1998) are based on misidentifications.

Cercospora trianthematis Chidd., Mycopathol. Mycol. Appl. 17: 80 (1962); as “trianthemae”. (Fig. 60)

Fig. 60
figure60

Cercospora trianthematis (K(M) IMI 83193, lectotype). A. Conidiophores fascicle. B. Conidia. Bar = 10 µm.

Synonym: Cercospora aizoacearum Bhartiya et al., Kavaka 25: 45 “1997” (1998) [holotype: India: Uttar Paradesh: Gorakhpur, on Trianthema portulacastrum [monogyna], Aizoaceae, Sep. 1998, H. D. Bhartiya (HCIO, s.n.); isotype: GPU 8073].

Literature: Crous & Braun (2003: 409), Kamal (2010: 14, 94).

Illustrations: Chiddarwar (1962: 78, plate II, figs 1213), Bhartiya et al. (1998: 45, fig. 2).

Description: Leaf spots amphigenous, circular to oval or irregular, scattered, 0.5–6 mm diam, dark brown to blackish. Casepituli amphigenous, mainly hypophyllous, punctiform, brown. Mycelium internal. Stromata substomatal, 15–45 µm diam, olivaceous to brown. Conidiophores in small to moderately large fascicles, 2–20, occasionally solitary, relatively dense to divergent, arising from stromata, through stomata, erect, straight, subcylindrical-conical to geniculate-sinuous, usually unbranched, 15–100 × 3–5 µm, 0–7-septate, dark olivaceous to brown, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal, intercalary or conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci thickened and darkened, 2.5–3 µm diam. Conidia solitary, shorter conidia obclavate to subcylindrical, longer ones acicular, straight to curved, (10−)30–125 × 2.5–4 µm, occasionally broader, (1−)3–11-septate, hyaline, thin-walled, smooth, apex pointed to subobtuse, base truncate to short obconically truncate, about 2–2.5 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202798): India: Maharashtra: Pune, Wanowri, Military Hospital, on Trianthema portulacastrum, 12 Oct. 1956, P. P. Chiddarwar 32 (K(M) IMI 83193). Isolectotypes: BPI 441988, HCIO.

Host range and distribution: On Trianthema portulacastrum, Aizoaceae, Asia (India, Maharashtra).

Note: A true Cercospora s. str. distinct from the C. apii s. lat. complex by having obclavate-cylindrical to acicular conidia. C. aizoacearum, described from India on the same host, is undoubtedly conspecific. Type material was not available. The conidia were described to be cylindrical, but the illustration shows cylindrical to somewhat obclavate shorter and acicular longer conidia, a range similar to C. trianthematis.

Altingiaceae

Key to cercosporoid species on Altingiaceae
  1. 1

    Leaf spots absent or almost so, sometimes with slight discolorations on the upper leaf surface; mycelium internal; superficial hyphae lacking; conidiophores fasciculate, 0–1-septate, 3–7 µm wide, conidiogenous loci unthickened, nor darkened; conidia 20–80 × 4–7.5 µm; on Liquidambar styraciflua, North America ………. Pseudocercospora tuberculans

    With distinct leaf spots; mycelium internal and often also external; conidiophores fasciculate and often also solitary, arising from superficial hyphae if present, aseptate, narrower, 2–5 µm wide; conidia much narrower, 2–4 µm ………. 2

  2. 2(1)

    Leaf spots small, 0.5–4 mm diam, dingy greenish grey, brown, grey, finally usually greyish white, with dark, often somewhat raised margin; caespituli mainly epiphyllous, conspicuous, punctiform to almost pustulate, dark brown to blackish; conidiophores pale to medium dark brown, paler towards the tip, wall somewhat thickened (to 1 µm), (10−)25–90 µm long; conidiogenous loci conspicuous, slightly thickened, at least around the outer rim, somewhat darkened-refarctive, 1–1.5 µm diam, in front view visible as minute circle; conidia to 150 µm long, subhyaline or very pale olivaceous, hila unthickened to slightly thickened, somewhat darkened-refractive; on Liquidambarstyraciflua, North America ………. Cercospora liquidambaris

    Leaf spots larger, usually 1–10 mm diam; caespituli less conspicuous, never pustulate; conidiophores paler, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, shorter, 5–35 µm long; conidiogenous loci inconspicuous, neither thickened not darkened; conidia shorter, to 100 µm, subhyaline or pale to medium olivaceous or olivaceous-brown, hila unthickened, not darkened on Liquidambar formosana and L. styraciflua, Asia and North America ………. P. liquidambaricola

Cercospora (s. lat.)

Cercospora liquidambaris Cooke & Ellis ex G.F. Atk., J. Elisha Mitchell Sci. Soc. 8: 48 (1892) (Fig. 61)

Fig. 61
figure61

Cercospora liquidambaris (CUP-A-002227(AL), lectotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophores. D. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 259).

Description: Leaf spots amphigenous, subcircular to angular-irregular, small, 0.5–4 mm diam, dingy greyish green, brown, to finally usually greyish white or white, sometimes somewhat raised, margin indefinite or usually darker, narrow. Caespituli amphigenous, usually epiphyllous, very conspicuous, punctiform to almost pustulate, scattered, dark brown to blackish. Mycelium internal and external; superficial hyphae lacking or almost so to well-developed, branched, septate, subhyaline to pale olivaceous or brownish, 1.5–4 µm wide. Stromata almost lacking or small to usually well-developed, substomatal or intraepidermal, 10–60 µm diam, brown. Conidiophores in small to moderately large fascicles, arising from substomatal hyphae or stromata, though stomata or erumpent, divergent to moderately dense, or solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical or attenuated towards the apex to usually geniculate-sinuous, unbranched, (10−)25–90(− 100) × 3–5 µm, (0−)1–8-septate, pale to medium brown, paler towards the tip, wall somewhat thickened, at least below, to 1 µm, smooth or almost so, proliferation sympodial and occasionally percurrent (conidiophores with enteroblastic rejuvenation leaving delicate annellations); conidiogenous cells integrated, terminal, 10–25 µm long, conidiogenous loci conspicuous, in front view visible as minute circle, 1–1.5 µm diam, at least rim slightly thickened, somewhat darkened or refractive. Conidia solitary, narrowly obclavate-cylindrical, filiform-subacicular, straight to curved, occasionally germinating with short to moderately long lateral outgrowths, 40–150 × (2−)2.5–4(− 4.5) µm, 3–12-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex acute to subobtuse, base usually short obconically truncate, sometimes truncate, 1–2 µm wide, hila unthickened or almost so to slightly thickened, somewhat darkened or refractive.

Lectotype (designated here, MycoBank, MBT202799): USA: Alabama: Auburn, on Liquidambar styraciflua, 14 Oct. 1891, G. F. Atkinson [CUP-A-002227(AL)].

Host range and distribution: On Liquidambar styraciflua, Altingiaceae, North America (Mexico; USA, Alabama, Delaware, Florida, Indiana, Louisiana, Maryland, Mississippi, Montana, Massachusetts, North Carolina, Texas).

Notes: The taxonomy and nomenclature of Cercospora liquidambaris have been totally confused. Chupp (1954) reduced C. liquidambaris Sawada, based on Taiwanese material causing a leaf spot disease of Liquibambar formosana, to synonymy with C. liquidambaris Cooke & Ellis ex G.F. Atk., which is, however, incorrect since the cercosporoid fungus on these hosts in Asia belongs in Pseudocercospora and is now correctly assigned to P. liquidambaricola (see below). The later species is also known from North America on Liquibambar styraciflua. Several collections from Mexico and the USA have been examined. The identity of the true C. liquidambaris is clarified by lectotypification. Atkinson (1892) cited a specimen collected in Alabama in 1891. A corresponding sample from Atkinson’s herbarium at CUP is designated as lectotype. The generic affinity of C. liquidambaris remains unclear. The conidiogenous loci and conidial hila are thickened and darkened. The conidia are cercospora-like, but pigmented as in the Passalora compex, which recently proved to be phylogenetically quite heterogeneous (S.I.R. Videira, unpubl.). Therefore, this species is tentatively retained in Cercospora s. lat. until phylogenetic data will be available.

Pseudocercospora

Pseudocercospora species on Altingiaceae

Pseudocercospora liquidambaricola (J.M. Yen) U. Braun, Schlechtendalia 5: 44 (2000). (Fig. 62)

Fig. 62
figure62

Pseudocercospora liquidambaricola (Hsieh & Goh 1990: 151, fig. 113). A. Conidiophores fascicles. B. Conidiophores arising from superficial hyphae. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Cercospora liquidambaricola J.M. Yen, Bull. Trimestriel Soc. Mycol. France 94: 52 (1978).

Synonyms: Cercospora liquidambaris Sawada, Rep. Gov. Agric. Res. Inst. Taiwan 85: 112 (1943), nom. inval. (Art. 39.1), and nom. illeg. (Art. 53.1), non C. liquidambaris Cooke & Ellis ex G.F. Atk., 1892 [type: Taiwan: Taipei, on Liquidambar formosana, 14 Oct. 1928, K. Sawada (NTU-PPE, hb. Sawada)].

Cercoseptoria liquidambaricola (J.M. Yen) J.M. Yen, Bull. Trimestriel Soc. Mycol. France 97: 92 (1981).

Pseudocercospora liquidambaris Goh & W.H. Hsieh, in Hsieh & Goh, Cercospora and similar fungi from Taiwan: 150 (1990) [type: Taiwan: Taipei, on Liquidambar formosana, 14 Oct. 1928, K. Sawada (NTU-PPE, hb. Sawada)].

Pseudocercospora neoliquidambaris C. Nakash. & Tak. Kobay., Mycoscience 43: 224 (2002), nom. illeg. [nom. superfl.] (Art. 52.1).

Literature: Chupp (1954: 259 p.p.), Hsieh & Goh (1990: 150), Guo & Hsieh (1995: 123–124), Guo et al. (1998: 138–140), Crous & Braun (2003: 253).

Illustrations: Yen (1978a: 53, fig. 3), Hsieh & Goh (1990: 151, fig. 113), Guo & Hsieh (1995: 125, fig. 110), Guo et al. (1998: 139, fig. 114), Kobayashi et al. (2002: 224, fig. 6).

Description: Leaf spots subcircular to angular-irregular, 1–10 mm diam or confluent and larger, angular spots often vein-limited, pale olivaceous, brown to dark brown or later greyish brown to greyish white, with dark border, brown to blackish, narrow, sometimes raised, with diffuse yellowish halo, occasionally somewhat zonate. Caespituli amphigenous, delicately punctiform to subeffuse, brown. Mycelium internal and external; superficial hyphae lacking or almost so to developed (mainly hypophyllous when developed), branched, 1.5–4 µm wide, subhyaline to pale olivaceous-brown, thin-walled, smooth. Stromata small to moderately large, 15–40 µm diam, rarely larger, substomatal to immersed, subglobose, brown. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, through stomata or erumpent, also solitary, arising from superficial hyphae if present, lateral, erect, straight, subcylindrical-conical to geniculate-sinuous, unbranched, 5–35 × 2–5 µm, 0–2-septate, subhyaline, pale olivaceous to olivaceous-brown, thin-walled, smooth to somewhat rough; conidiophores reduced to conidiogenous cells or integrated, terminal, 5–25 µm long, conidiogenous loci inconspicuous or visible as a truncate tip, but always unthickened and not darkened. Conidia solitary, cylindrical to obclavate-cylindrical, straight to curved, occasionally sigmoid, 20–100 × 2–4 µm, 2–10-septate, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth, apex obtuse to subacute, base short obconically truncate, sometimes truncate, 1–2 µm wide, hila unthickened, not darkened

Neotype (designated here, MycoBank, MBT202800): Taiwan: Taipei, on Liquidambar formosana, 14 Oct. 1928, K. Sawada (NTU-PPE, hb. Sawada).

Host range and distribution: On Liquidambar (formosana, styraciflua), Altingiaceae, Asia (China, Japan, Taiwan), North America (Mexico; USA, Alabama, Florida, Louisiana, Texas).

Notes: The holotype material of Cercospora liquidambaricola (Taiwan: Taichung, on Liquidambar formosana, 29 Oct. 1971, J.-M. Yen 71277 bis) could not be traced in PC or UC. Therefore, we designate a neotype here. Collections of Pseudocercospora on Liquidambar formosana in Asia and L. styraciflua in North America are morphologically very similar. If they are truly conspecific or if two morphologically very close, but geographically and genetically distinct species are involved can only be clarified on the base of molecular sequence analyses. Loropetalum chinense (Hamamelidaceae) was reported as host of this species from China (Guo & Hsieh 1995; Guo et al. 1998). The identity of Pseudocercospora on this host is unclear and needs to be confirmed. Chupp (1954), who proposed the original synonymy, found material from the USA and Taiwan to represent the same fungus. However, Hsieh & Goh (1990) did not examine American material. Samples from Taiwan (Taichung, 8 Aug. 1945, K. Sawda, BPI 437752, 437753) are probably topotypes. Guo & Hsieh (1995) confirmed the synonymy of C. liquidambaricola. They mentioned that they had examined Yen’s type material, but this material was not cited under “material examined”. The material concerned is not preserved at PC. The complicated nomenclature and taxonomy of this species has been discussed by Braun (2000a: 44) who emphasized that identity and application of the name C. liquidambaris can only be clarified by lectotypification. Chupp (1954) interpreted this name in the sense of P. liquidambaricola, which is clearly distinct from P. tuberculans by much narrower conidia, but his interpretation was based on a specimen collected by Geo. V. Nash in 1895 (USA, Florida, Lake City, Plants of Florida No. 2231, 11–19 Jul. 1895, BPI 437755, 437761, 437812), which is nontype material. Atkinson (1892) cited a specimen collected in Alabama in 1891, which can be used as lectotype. There is a single specimen in Atkinson’s herbarium (now CUP) which undoubtedly refers to this collection.

Pseudocercospora tuberculans (Ellis & Everh.) U. Braun, Schlechtendalia 2: 27 (1999). (Fig. 63)

Fig. 63
figure63

Pseudocercospora tuberculans (NY 2425377, lectotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora tuberculans Ellis & Everh., J. Mycol. 4: 115 (1888).

Literature: Saccardo (1892: 652), Chupp (1954: 259), Crous & Braun (2003: 412).

Exsiccatae: Ellis & Everh., Fungi Columb. 168. Ellis & Everh., N. Amer. Fungi 2292.

Description: Leaf spots absent or almost so, sometimes with slight discolorations on the upper leaf surface. Caespituli hypophyllous, punctiform, on small, brown, tubercle-like swellings, 0.5–1 mm diam, sometimes effuse between such swellings. Mycelium internal. Stromata almost absent to well-developed, 10–80 µm diam, dark brown to blackish, substomatal to immersed, composed of swollen hyphal cells, 3–8 µm diam. Conidiophores in small to large fascicles, loose to usually dense, subcylindrical to conical, straight to slightly geniculate-sinuous, unbranched, 10–35 × 3–7 µm, rarely longer, 0–1-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci inconspicuous or visible as truncate tips, always unthickened and not darkened. Conidia solitary, cylindrical to somewhat obclavate-cylindrical, straight to slightly curved, 20–80 × 4–6.5 µm, 1–6-septate, pale olivaceous to brownish, thin-walled, smooth, apex obtuse, base subtruncate to short obconically truncate, rarely long obconically truncate, 2–3 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202801): USA: Mississippi: Oktibbeha County, Starkville, on Liquidambar styraciflua, 27 June 1888, S. M. Tracy (NY 2425377). Isotypes: BPI 442929, 442032, 1102025; Ellis & Everh., N. Amer. Fungi 2292, e.g. BPI 442930, MICH 15380, NY 2425378–2425383. Topotypes: June 1890–1893 (BPI 442031, 442033, 442034; CUP — A(S.M.T.[03]); HBG, Ellis & Everh., Fungi Columb. 168).

Host range and distribution: On Liquidambar styraciflua, Altingiaceae, North America (USA, Florida, Louisiana, Missouri, Mississippi).

Notes: Records of this species from China on Liquidambar formosana (Keissler & Lohwag 1937, Tai 1979) are doubtful, unproven and not included in Guo & Hsieh (1995) and Guo et al. (1998). They refer to Chinese material on dead wood of Liquidambar formosana distributed as Cercospora tuberculans in Petr., Crypt. Exs. 3391 [Hunan, Changcha, Nov., C. Keissler & H. Handel-Mazzetti (W)]. The identity of this wood-inhabiting fungus is not quite clear, but it undoubtedly does not pertain to C. tuberculans.

Amaranthaceae s. str.

(The family Chenopodiaceae is phylogenetically close to Amaranthaceae and currently proposed to be included in the latter family (s. lat.), but we prefer to maintain the Chenopodiaceae as separate family, at least tentatively.)

Cercospora

Key to Cercospora species on Amaranthaceae
  1. 1

    Conidia cylindrical or obclavate-cylindrical ………. 2

    Conidia consistently acicular or at least longer conidia acicular and only shorter ones somewhat obclavate-cylindrical ………. 5

  2. 2(1)

    Stromata lacking or almost so; conidiophores short, 5–30 × 3–5 µm, 0(− 1)-septate; conidia short, 25–35 × 2.5–3.5 µm, 0–5-septate; on Celosia sp ………. C. celosiigena

    Stromata developed, 10–60 µm diam; conidiophores longer, 10–150 µm, 0–8-septate; and/or conidia longer, 15–150 µm, pluriseptate ………. 3

  3. 3(2)

    Conidiogenous loci cercospora-like, i.e. distinctly thickened and darkened, in front view visible as small dark circle with minute central pore; conidiophores distinctly brown; on Celosia sp ………. C. gorakhanathii

    Conidiogenous loci not cercospora-like, i.e. unthickened to somewhat thickened, but not darkened, at most somewhat refractive, formed on characteristic, bulging, convex tips and shoulders caused by sympodial proliferation; conidiophores hyaline, only pale olivaceous at the base or pale olivaceous-brown, tips often hyaline or subhyaline ………. 4

  4. 4(3)

    Conidiophores hyaline or only pale olivaceous at the base; conidia obclavate-cylindrical, 20–80 × 3.5–7 µm; on Achyranthes spp ………. Cercosporella pseudachyranthis (see “Doubtful, excluded and insuffiently known species of Cercospora”)

    Conidiophores pale olivaceous-brown throughout or tips paler, hyaline or subhyaline; conidia narrowly cylindrical to subacicular, 15–90 × 2–4.5 µm; on Gomphrena spp ………. C. pretoriensis (see “Doubtful, excluded and insuffiently known species of Cercospora”)

  5. 5(1)

    Leaf spots lacking or almost so; conidia acicular to obclavate-cylindrical, broad, 35–195 × (3−)4–6.5(− 8) µm; on Achyranthes spp ………. C. achyranthis

    Leaf spots developed, distinct; conidia consistently acicular and/or narrower, 1.5–5 µm ………. 6

  6. 6(5)

    Conidia narrowly cylindrical to subacicular, apex obtuse, not distinctly pointed; on Gomphrena spp., South Africa ………. C. pretoriensis

    At least longer conidia distinctly acicular with truncate base and acute or subacute apex; on other hosts ………. 7

  7. 7(6)

    Conidia consistently acicular, base truncate, various morphologically barely distinguishable species (see Tabular key to Cercospora species on Acanthaceae according to host genera)

    Conidia acicular to obclavate-cylindrical, at least shorter conidia obclavate-cylindrical, with obconically truncate base ………. 8

  8. 8(7)

    Stromata well-developed, large, 20–80 µm diam; conidiophores 10–80 µm long; conidial base 1–2 µm wide; on Alternanthera spp ………. C. alternantherae

    Stromata lacking or 10–50 µm diam; conidiophores to 250 µm long; conidial base 1.5–4 µm wide; on Amaranthus spp ………. C. brachiata

Tabular key to Cercospora species on Acanthaceae according to host genera

Achyranthes

  1. 1

    Conidiophores colourless or only olivaceous at the base; conidiogenous loci conspicuous, but not cercospora-like (not darkened, only refractive, characteristically bulging, cercosporella-like) ………. Cercosporella pseudachyranthis (see “Doubtful, excluded and insufficiently known species of Cercospora”)

    Conidiophores brown, distinctly pigmented throughout; conidiogenous loci distinctly thickened and darkened, Cercospora type ………. 2

  2. 2(1)

    Leaf spots lacking or almost so; conidiophores to 450 µm long; conidia acicular to obclavate-cylindrical, 35–195 × (3−)4–6.5(− 8) µm ………. C. achyranthis

    Leaf spots distinct, well-developed; conidiophores to 250 µm long; conidia usually consistently acicular, much narrower, 25–240 × 2–5 µm ………. C. achyranthina

  • Aerva

    A single species ………. C. aervae-lanatae

  • Alternanthera

    A single species ………. C. alternantherae

  • Amaranthus

    A single species ………. C. brachiata

  • Celosia

    1. 1

      Conidiophores short, 5–30 × 3–5 µm, 0(− 1)-septate; conidia short, 25–35 × 2.5–3.5 µm, cylindrical to obclavate-cylindrical, 0–5-septate, hila 1–2 µm wide ………. C. celosiigena

      Conidiophores much longer, 20–220 µm, 0–8-septate; conidia much longer, 15–150 µm, (0−)1–14-septate, hila 1.5–3 µm wide ………. 2

    2. 2(1)

      Stromata lacking or almost so; conidia acicular ………. C. celosiae

      Stromata developed, 10–40 µm diam; conidia cylindrical or obclavate-cylindrical ………. C. gorakhanathii

  • Cyathula

    A single species ………. C. apii s. lat. (C. cf. maloti sensu Groenewald et al. 2013)

  • Digera

    A single species ………. C. achyranthina

  • Gomphrena

    1 Stromata lacking or small, 10–35 µm diam; conidia acicular, 30–450 µm long, apex pointed; mainly on Gomphrena globosa, Northeast Africa, Asia, North America, West Indies ………. C. gomphrenae

    Stromata 15–60 µm diam; conidiogenous loci not cercospora-like (neither distinctly thickened, nor darkened), conidia narrowly cylindrical to subacicular, apex obtuse; on Gomphrena spp., South Africa ………. C. pretoriensis (see “Doubtful, excluded and insuffiently known species of Cercospora”)

  • Pupalia

    A single species ………. C. achyranthina

Cercospora species on Amaranthaceae

Cercospora achyranthina Thirum. & Chupp, Mycologia 40: 352 (1948). (Fig. 64)

Fig. 64
figure64

Cercospora achyranthina (BPI 4323844, holotype). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 29), Vasudeva (1963: 31), Ibrahim & El Nur Elamin (1974), Ellis (1976: 244), Crous & Braun (2001: 328), Crous & Braun (2003: 42), Guo et al. (2005: 27), Kamal (2010: 12), Braun & Urtiaga (2013: 177).

Illustrations: Ellis (1976: 243, fig. 183 B), Guo et al. (2005: 27, fig. 9).

Description: Leaf spots amphigenous, circular to angular-irregular, 0.5–6.5 mm diam, occasionally confluent and larger, to 10 mm diam, brown to greyish brown, finally sometimes dull grey or greyish white, margin reddish brown to purple-violet, finally very dark, sometimes with diffuse darker halo, often purplish. Caespituli amphigenous, scattered, finely punctiform, dark. Mycelium internal. Stromata almost lacking or small, 10–25 µm diam, substomatal to immersed, brown. Conidiophores in fascicles, 2–10, divergent, arising from swollen hyphal cells or stromata, through stomata or erumpent, erect, straight to curved, geniculate-sinuous, above all in the upper half, unbranched, 15–250 × 3–8 µm, (0−)1–10-septate, pale olivaceous-brown or brownish, wall somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, 10–40 µm long, conidiogenous loci thickened and darkened, 2–3.5 µm diam. Conidia solitary, acicular, shorter conidia occasionally slightly obclavate-cylindrical, (25−)35–150(− 240) × 2–5 µm, usually 3–10-septate, hyaline, thin-walled, smooth, apex acute or subobtuse, base truncate, occasionally slightly obconically truncate, 2–3.5 µm wide, hila thickened and darkened.

Holotype: India: Karnataka: Bangalore, on Achyranthes aspera, 20 Aug. 1944, M. J. Thirumalachar (BPI 4323844). Topotype: 2 Sep. 1945 (CUP 37201).

Host range and distribution: On Achyranthes (aspera, Achyranthes sp.), Digera muricata, Pupalia lappacea [atropurpurea]. Amaranthaceae, Africa (Sudan, Tanzania, Zimbabwe), Asia (China; India, Bihar, Karnataka, Andra Pradesh, Madhya Pradesh, Maharashtra, New Delhi, West Bengal; Myanmar, Pakistan, Philippines), South America (Venezuela).

Notes: This species is part of the Cercospora apii s. lat. complex. Thirumalachar & Chupp (1948) cited a single collection from 1944 as type. Chupp (1954) erroneously mentioned a collection from 2 Sep. 1945 as type material. This specimen, which can be considered topotype material, is maintained and deposited as CUP 37201.

Cercospora achyranthis Syd. & P. Syd., Ann. Mycol. 7: 171 (1909). (Fig. 65)

Fig. 65
figure65

Cercospora achyranthis (LEP, isolectotype). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Saccardo (1913: 1429), Vassiljevsky & Karakulin (1937: 222), Chupp (1954: 30), Vasudeva (1963: 31), Katsuki (1965: 8), Shin & Kim (2001: 24), Braun & Crous (2003: 42), Guo et al. (2005: 28–29), Kamal (2010: 12).

Illustrations: Shin & Kim (2001: 25, fig. 1), Guo et al. (2005: 28, fig. 10).

Exsiccatae: Cif., Mycofl. Doming. Exs. 147. Syd., Fungi Exot. Exs. 546.

Description: Leaf spots lacking or almost so, indistinct or only diffuse discolorations, greenish to brownish, finally sometimes vein-limited, greyish, margin indefinite. Caespituli hypophyllous, effuse, floccose-velutinuous, dark, greyish to dark grey or brownish. Mycelium internal; hyphae branched, septate, somewhat pigmented. Stromata lacking or almost so. Conidiophores in small, loose fascicles, 2–12, arising from substomatal swollen hyphal cells, through stomata, erect, straight to usually distinctly geniculate or geniculate-sinuous, often strongly geniculate, unbranched, 40–450 × 3.5–8 µm, pluriseptate, pale to medium dark olivaceous or olivaceous throughout or paler towards the tip, wall thin to somewhat thickened, smooth; conidiogenous cells integrated, terminal and intercalary, about 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, 1.5–2.5 µm diam. Conidia solitary, acicular to obclavate-cylindrical, straight to curved, 35–195 × (3−)4–6.5(− 8) µm, 3–18-septate, hyaline or almost so, thin-walled, smooth, apex acute to subobtuse, base truncate to short obconically truncate, 1.5–4 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202802): Japan: Saitama Prefecture: Ome, Musashi, on Achyranthes bidentata, 22 Sep. 1905, I. Miyake (S-F23053). Isolectotypes: LEP.

Host range and distribution.: On Achyranthes (aspera [indica], bidentata [japonica]), Amaranthaceae, Asia (China; India, Andhra Pradesh, Maharashtra, Rajasthan; Japan, Korea, Pakistan, Taiwan), West Indies (Dominican Republ., Puerto Rico).

Note: A true Cercospora s. str. distinct from C. apii s. lat., including C. achyranthina, by having acicular to obclavate-cylindrical conidia with obconically truncate base. Obconically truncate conidial bases are not confined to shorter, young conidia, they are also evident in longer, fully developed conidia. In addition, different from C. achyranthina by lacking or indistinct leaf spots, effuse caepituli and somewhat wider conidia.

Cercospora aervae-lanatae Raghu Ram & Mallaiah, Mycol. Res. 100: 296 (1996); as “aerva-lanatae”. (Fig. 66)

Fig. 66
figure66

Cercospora aervae-lanatae (K(M) IMI 351224, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonym: Cercospora aervae R.K. Srivast. et al., Indian Phytopathol. 54: 102 (2001); as “aeruae” [holotype: India: Uttar Pradesh: Gorakhpur, on Aerva sanguinolenta [scandens], Nov. 1990, R. K. Srivastava 123 (GPU 1398). Isotype: HCIO 30878.].

Literature: Crous & Braun (2003: 47), Kamal (2003: 13).

Illustrations: Raghu Ram & Mallaiah (1996: 296, fig. 2), Srivastava et al. (2001: 102, fig. 1).

Description: Leaf spots amphigenous, necrotic, scattered, subcircular to angular-irregular, 2–9(− 10) mm diam, greyish, sometimes with reddish or purplish margin. Caespituli epiphyllous. Mycelium immersed; hyphae branched, septate, subhyaline. Stromata well-developed, gobose, 20–40 µm diam, brown to black. Conidiophores in loose fascicles, to 15, arising from stromata, through stomata, erect, subcylindrical to 1–4 times geniculate, unbranched, about 45–230 × 3.5–5.5 µm, 2–11-septate, pale olivaceous to brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci thickened and darkened, 2–3.5 µm diam. Conidia solitary, acicular, straight to curved, 55–160 × 1.5–4 µm, 5–15-septate, hyaline, thin-walled, smooth, apex pointed, base truncate, 1.5–3 µm wide, hila thickened and darkened.

Holotype: India: Andhra Pradesh: Nagarjuna Nagar, University Campus, on Aerva lanata, Nov. 1991, M. Raghu Ram (K(M) IMI 351224).

Host range and distribution: On Aerva (lanata, sanguinolenta [scandens]), Amaranthaceae, Asia (India, Andhra Pradesh, Uttar Pradesh).

Note: This species is morphologically assignable to the Cercospora apii s. lat. complex.

Cercospora alternantherae Ellis & Langl., J. Mycol. 6: 36 (1890). (Fig. 67)

Fig. 67
figure67

Cercospora alternantherae (NY 830163, lectotype). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Saccardo (1892: 637), Chupp (1954: 30–31), Crous & Braun (2003: 53), Kamal (2010: 15).

Illustration: Chupp (1954: 31, fig. 7).

Description: Leaf spots amphigenous, circular or subcircular, 0.5–3 mm diam, pale greenish to dingy grey, margin brownish. Caespituli amphigenous, punctiform, scattered, dark brown to blackish. Mycelium internal. Stromata substomatal to immersed, large, 20–80 µm diam, brown. Conidiophores in well-developed, mostly rather large fascicles, divergent to dense, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical-conical to oblong, geniculate-sinuous, unbranched, 10–80 × 3–6 µm, 1–4-septate, subhyaline, pale olivaceous to brown, paler towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, (1−)1.5–2(− 2.5) µm diam. Conidia solitary, obclavate-cylindrical, acicular, straight to curved, 20–125 × 2–4 µm, 0–10-septate, hyaline, thin-walled, smooth, apex pointed or subobtuse, base truncate to obconically truncate, 1–2 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202806): USA: Louisiana: St. Martinsville, on Alternanthera achyrantha, 18 Jul. 1888, A. B. Langlois 1430 (NY 830163). Isolectotype: BPI 432464.

Host range and distribution: On Alternanthera (ficoidea, halimifolia, crucis [portoricensis], pungens [achyrantha, repens]), Amaranthaceae, North America (USA, Louisiana, Texas), South America (Brazil, Venezuela), West Indies (Cuba, Jamaica, Puerto Rico, Virgin Islands).

Notes. This is a true Cercospora s. str. close to C. apii s. lat., but distinct by having large stromata (20–80 µm diam) with numerous, densely arranged conidiophores, smaller conidiogenous loci (1–2 µm wide), and obclavate-cylindrical to acicular conidia. Indian records of this species from Uttar Pradesh and West Bengal on A. sessilis (Kamal 2010) are doubtful and in need of revision and confirmation. They might rather pertain to the illegitimate species Cercospora sessilis or to the invalid C. alternantherina, both described from India on A. sessilis.

Cercospora apii s. lat. (C. cf. malloti sensu Groenewald et al. 2013: 157).

Notes: Nguanhom et al. (2015) examined Cercospora species from northern Thailand by means of molecular methods. A plurivorous Cercospora species referred to as Cercospora cf. malloti in Groenewald et al. (2013) turned out to be the most common taxon found during the course of this study. The collections in the clade concerned belong to C. apii s. lat., i.e. they are characterised by having consistently acicular conidia. One of the specimens involved was collected on Cyathula prostrata. For further comments, see notes under Cercospora asystasiana.

Cercospora brachiata Ellis & Everh., J. Mycol. 4: 5 (1888). (Fig. 68)

Fig. 68
figure68

Cercospora brachiata (NY 2408319, lectotype). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonyms: Cercospora acnidae Ellis & Everh., Proc. Acad. Nat. Sci. Philadelphia 43: 89 (1891) [lectotype (designated here, MycoBank, MBT202807): USA: Delaware: Wilmington, on Amaranthus cannabinus, 30 Sep. 1889, A. Commons 1011 (NY 2408316); isolectotypes: NY 2408317, 2408318].

Cercospora amaranthi Lobik, Bolezni Rast. 17: 193 (1928) [holotype: Russia: Stavropol Krai: Pyatigorsk, station of Yessentuki, garden, on Amaranthus retroflexus, 25 Sep. 1925, A. I. Lobik (not preserved)].

Literature: Saccardo (1892: 637), Vassiljevsky & Karakulin (1937: 222), Chupp (1954: 30–31), Katsuki (1965: 8), Braun & Mel’nik (1997: 43–44), Crous & Braun (2003: 43, 86), Guo et al. (2005: 29–30), Kamal (2010: 25).

Illustrations: Tai (1948: 37, fig. 2), Guo et al. (2005: 29, fig. 11), Pirnia et al. (2010: 185, fig. 1).

Exsiccatae: Ellis & Everh., N. Amer. Fung. 2582.

Description: Leaf spots amphigenous, circular, subcircular to somewhat angular-irregular, 0.5–12 mm diam, at first yellowish brown, later brown, reddish or dark brown, or finally with a tan, grey to greyish white centre surrounded by a brown, reddish to purplish brown or almost blackish margin. Caespituli amphigenous, mostly hypophyllous, punctiform scattered, dark. Mycelium internal. Stromata almost lacking or small to moderately large, substomatal to immersed, subglobose to somewhat irregular, 10–50 µm diam, olivaceous-brown to brown, cells 2–6 µm diam. Conidiophores fasciculate, 2–12(− 20), rarely solitary, arising from swollen hyphal cells or stromata, emerging through stomata or erumpent, erect, straight, subcylindrical to usually distinctly geniculate or geniculate-sinuous, above all in the upper half, unbranched or tips occasionally furcate, 20–250 × 3–6.5 µm, usually 1–8-septate, very long conidiophores sometimes with additional septa, pale to medium dark brown or olivaceous-brown throughout or paler towards the tip, wall thin to slightly thickened, smooth; conidiogenous cells integrated, terminal and intercalary, 10–40(− 70) µm long, conidiogenous loci conspicuous, thickened and darkened, 1.5–3.5 µm diam. Conidia solitary, acicular, shorter conidia may also be narrowly obclavate-cylindrical, straight to curved, 25–250 × 1.5–5(–6) µm, 1–20-septate, hyaline, thin-walled, smooth, apex acute or subacute, sometimes subobtuse, base truncate or obconically truncate in shorter conidia, 1.5–4 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202808): USA: Delaware: Faulkland, on Amaranthus retroflexus, 18 Aug. 1887, A. Commons 626 (NY 2408319). Isolectotypes: NY 24083120, 2408321.

Host range and distribution: On Amaranthus (albus, blitum, blitoides, cannabinus, caudatus, crassipes, cruentus [paniculatus, hybridus subsp. cruentus], dubius, hybridus, polygamus, retroflexus, spinosus, tamariscinus, tricolor [gangeticus], tuberculatus, viridis [blitum subsp. emarginatus], Amarathus sp.), Amaranthaceae, Africa (Kenya, South Africa, Uganda), Asia (Brunei, China; India, Andhra Pradesh, Delhi, Orissa, Maharashtra, Uttar Pradesh, West Bengal; Indonesia, Iran, Pakistan), Europe (Germany, Russia, Ukraine), Central and South America (Brazil, Ecuador, Panama, Venezuela), North America (USA, Delaware, Florida, Illinois, Nebraska, Texas, Wisconsin), West Indies (Barbados, Cuba, Dominican Republic, Haiti, Puerto Rico, Trinidad and Tobago, Virgin Islands).

Notes: This species belongs to the Cercospora apii s. lat. complex. Cercospora brachiata is morphologically rather variable, above all with regard to the length of conidiophores as well as length and width of conidia. Many host species and a wide distribution range covering different continents are involved. Thus, it remains unclear if all collections pertain to a single species or if we have to do with a complex of cryptic species. Cercospora acnidae is morphologically barely distinct from C. brachiata. Its introduction and recognition in Chupp (1954) were undoubtedly influenced by assumed host range differences. The hosts of C. acnidae were previously assigned to the genus Acnidia, now a synonym of Amaranthus usually treated as subgenus, which reflects the close affinity of former Acnida and Amaranthus species. A careful search for type material of Cercospora amaranthi in LE failed. Type material of this species is probably not maintained. Records of C. brachiata on Achyranthes bidentata [japonica] (Crous & Braun 2003) are doubtful and belong probably to C. achyranthina. Some newer records not yet included in Crous & Braun (2003) refer to Germany (Jage & Braun 2004), Indonesia (Shivas et al. 1996) and Iran (Pirnia et al. 2010, Hesami et al. 2011).

Ellis & Everh., N. Amer. Fung. 2582 (BPI 432395, FH, GZU, NY, PH and numerous other herbaria) is authentic material (former syntypes) from the type locality, but collected in 1890.

Cercospora celosiae Syd., Ann. Mycol. 27: 430 (1929). (Fig. 69)

Fig. 69
figure69

Cercospora celosiae (BPI 434404, holotype). A. Conidiophore fascicle. B. Conidiophore tip. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 32), Vasudeva (1963: 76), Katsuki (1965: 9), Ibrahim & El Nur Elamin (1974), Ellis (1976: 244), Hsieh & Goh (1990: 15), Crous & Braun (2003: 113), Guo et al. (2005: 30–31), Kamal (2010: 31).

Illustrations: Ellis (1976: 243, fig. 183 C), Hsieh & Goh (1990: 17, fig. 6), Guo et al. (2005: 31, fig. 12).

Description: Leaf spots amphigenous, circular or subcircular, 1–12 mm diam, occasionally larger, tan to pale brown, margin darker, occasionally somewhat raised, sometimes causing shot-hole symptoms. Caespituli usually hypophyllous, delicate, dark. Mycelium internal. Stromata lacking or small, brown, substomatal. Conidiophores in fascicles, 2–15, divergent, arising from internal hyphae or small hyphal aggregations, through stomata, erect, straight, usually unbranched, geniculate-sinuous, 20–220 × 3–6 µm, 0–6-septate, pale to medium brown, paler and narrower towards the tip, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened, 1.5–3 µm diam. Conidia solitary, acicular, straight to curved, 25–150 × 2–4(− 4.5) µm, hyaline, 2–12-septate, apex acute, base truncate to somewhat obconically truncate, 1.5–3 µm wide, hila thickened and darkened.

Holotype: China: Hubei: Wang-Chia-Shau, on Celosia argentea, 4 Aug. 1928, T F Yu (BPI 434404).

Host range and distribution: On Celosia (argentea [argentea var. cristata, cristata, plumosa], trigyna [laxa], Celosia sp.), Amaranthaceae, Africa (Nigeria, Sudan, Uganda,), Asia (Bangladesh, Brunei, Cambodia, China, India, Indonesia, Malaysia, Myanmar, Pakistan, Papua New Guinea, South Korea, Sri Lanka, Taiwan, Thailand), Central and South America (Brazil, Panama, Venezuela), North America (Mexico; USA, Alabama, Florida, Oklahoma), West Indies (Cuba).

Notes: This species is characterised by having colourless acicular conidia and thickened, darkened conidiogenous loci and hila, i.e. it belongs to the Cercospora apii s. lat. complex. Japanese records of this species (Katsuki 1965) are wrong and refer to Pseudocercospora celosiarum (confirmed by Ch. Nakashima).

Cercospora celosiigena U. Braun & Bagyan., sp. nov.

MycoBank MB814563 (Fig. 70)

Fig. 70
figure70

Cercospora celosiigena (HAL 2898 F, holotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Literature: Bagyanarayana et al. (1991: 324).

Illustration: Bagyanarayana et al. (1991: 321, fig. 6).

Diagnosis: Differs from C. celosiae in having much shorter conidiophores, 5–30 × 3–5 µm, 0(− 1)-septate, and short, narrowly obclavate-cylindrical conidia, 25–35 × 2.5–3.5 µm, only 0–5-septate,

Description: Leaf spots amphigenous, subcircular to angular-irregular, 1–3 mm diam, somewhat raised, greyish brown to greyish white, with a narrow purplish margin. Caespituli hypophyllous, scattered, fine, dark brown. Mycelium internal. Stromata lacking or small, formed by a few swollen hyphal cells, brown, substomatal. Conidiophores in small, divergent to dense fascicles, arising from substomatal hyphae or small stromatic hyphal aggregations, through stomata, erect, subcylindrial, conical, straight to somewhat curved or slightly geniculate-sinuous, unbranched, 5–30 × 3–5 µm, 0(−1)-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiophores usually aseptate, i.e. reduced to conidiogenous cells, conidiogenous cells occasionally integrated, terminal, 5–25 µm long, conidiogenous loci conspicuous, thickened and darkened, 1–2 µm diam. Conidia solitary, narrowly obclavate-cylindrical, straight to slightly curved, 25–35 × 2.5–3.5 µm, 0–5-septate, hyaline, thin-walled, smooth, apex acute to subobtuse, base truncate in cylindrical conidia to short obconically truncate in obclavate ones, 1–2 µm wide, hila slightly thickened and darkened.

Holotype: India: Telangana: Hyderabad, on Celosia argentea, Amaranthaceae, Oct. 1990, G. Bagyanarayana & P. Jagadeeswar (HAL 2898 F).

Host range and distribution: Only known from the type collection.

Notes: Bagyanarayana et al. (1991) identified the type collection of this species as Cercospora celosiae which they considered a morphologically rather variable fungus. However, C. celosiae belongs to the C. apii s. lat. complex. The conidia are consistently acicular, and the conidiophores are much longer and septate.

Cercospora gomphrenae W.W. Ray, Mycologia 36: 172 (1944). (Fig. 71)

Fig. 71
figure71

Cercospora gomphrenae (CUP 33132, holotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Literature: Chupp (1954: 34), Ibrahim & El Nur Elamin (1974), Crous & Braun (2003: 201), Shin & Kim (2001: 67–69), Guo et al. (2005: 31), Kamal (2010: 47).

Illustrations: Shin & Kim (2001: 68, fig. 25), Guo et al. (2005: 32, fig. 13).

Description: Leaf spots amphigenous, circular to somewhat angular-irregular, 1–10 mm diam, at first brownish, greyish brown, later dingy grey to greyish white, margin darker, brown, yellowish brown, reddish brown to purplish brown. Caespituli amphigenous, punctiform, dark brown, scattered. Mycelium internal; hyphae branched, septate, subhyaline or pale, 2–3.5 µm wide. Stromata almost lacking or 10–35 µm diam, substomatal to immersed, brown, cells 2–5 µm diam. Conidiophores in small, divergent fascicles, 3–15, arising from internal swollen hyphae or stromata, through stomata or erumpent, erect, straight, subcylindrical to distinctly geniculate-sinuous, unbranched, 30–300 × 3–7 µm, usually 3–10-septate, pale to medium brown, paler towards the tip, thin-walled, smooth; conidiogenous cells intergrated, terminal and intercalary, 10–30 µm long, conidiogenous loci conspicuously thickened and darkened, 2–3.5 µm diam. Conidia solitary, acicular, straight to curved, 30–300(− 450) × 2–5 µm, 3–20-septate, hyaline, thin-walled, smooth, apex acute to subobtuse, base truncate, 2–4 µm wide, hila thickened and darkened.

Holotype: USA: Oklahoma: Stillwater, Ray’s yard, on Gomphrena globosa, 18 Aug. 1942, W. W. Ray (CUP 33132). Isotype: K(M) IMI 168993. Topotype: from 8 Aug. 1942 (CUP 39900).

Host range and ditribution: On Gomphrena (globosa, serrata [decumbens]), Amaranthaceae, Africa (Sudan), Asia (China; India, Andhra Pradesh; Korea), North America (USA, Georgia, Oklahoma, Texas), West Indies (Cuba).

Notes: This species is part of the Cercospora apii s. lat. complex. Due to confusion with the name Cercospora gomphrenaea Sawada, several records of C. gomphrenae W.W. Ray are undoubtedly wrong or doubtful, e.g. those from Iran, Nepal and Taiwan (Crous & Braun 2003).

Cercospora gorakhanathii A.N. Rai & Kamal, Trans. Brit. Mycol. Soc. 89: 124 (1987). (Fig. 72)

Fig. 72
figure72

Cercospora gorakhanathii (K(M) IMI 259305, holotype). A. Conidiophores fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonym: Cercospora celosiicola Bhartiya et al., Indian Phytopathol. 56: 271 (2003) [holotype: India: Uttar Pradesh: Basti, on Celosia sp. (as “coronata”), Nov. 1997, H. D. Bhartiya (HCIO 42689); isotype: GPU 8026].

Literature: Kamal (2010: 31).

Illustrations: Rai & Kamal (1987: 125, fig. 1), Bhartiya et al. (2003: 271, fig. 2).

Description: Leaf spots amphigenous, circular to angular-irregular, sometimes vein-linited, 2–8 mm diam or confluent and larger, light brown to dingy grey. Caespituli amphigenous, finely punctiform, effuse, dark dingy olivaceous to brown. Mycelium internal. Stromata developed, substomatal to immersed, 10–40 µm diam, medium to dark olivaceous. Conidiophores fasciculate, divergent, 2–12 or occasionally solitary, arising from stromata, through stomata or erumpent, erect, straight, subcylindrical to geniculate-sinuous, usually unbranched, rarely branched, 20–150 × 3–6 µm, 1–8-septate, pale, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or occasionally intercalary, conidiogenous loci conspicuous, 1.5–2 µm diam, thickened and darkened. Conidia solitary, cylindrical, somewhat cylindrical-obclavate to subacicular, straight to curved, 15–150 × 1.5–4.5 µm, (0−)1–14-septate, hyaline, thin-walled, smooth, apex subacute or subobtuse, base truncate to somewhat obconically truncate, 1.5–3 µm wide, hila thickened and darkened.

Holotype: India: UttarPradesh: Gorakhpur, University, botanical garden, on Celosia sp., Feb. 1981, A. N. Rai KR 536 (K(M) IMI 259305).

Host range and distribution: On Celosia sp., Amaranthaceae, Asia (India, Uttar Pradesh).

Note: A true Cercospora s. str. well characterised by having cylindrical to somewhat obclavate or subacicular conidia.

Doubtful, excluded and insufficiently known species

Cercospora alternantherina S. Narayan et al., in Rao et al., Sugarcane Pathology 1: Fungal Diseases: 62 (1999), nom. inval. (Art. 39.1).

Literature: Crous & Braun (2003: 53), Kamal (2010: 15).

Illustration: Rao et al. (1999: 63, fig. 3).

Description: Leaf spots amphigenous, circular to subcircular, 0.5–4 mm diam, olivaceous to olivaceous-brown on the upper leaf surface, pale olivaceous with brown margin below. Caespituli amphigenous, effuse. Mycelium internal; hyphae branched, septate, subhyaline to light olivaceous, about 2.5 µm wide. Stromata well-developed, substomatal, about 25 µm diam, pale olivaceous to olivaceous. Conidiophores solitary or in fascicles of 2–6, arising from stromata, through stomata, about 15–130 × 2–4.5 µm, cylindrical, straight to slightly curved, unbranched, 1–4 times geniculate, pale to dark olivaceous, smooth; conidiogenous cells integrated, terminal and intercalary, with conspicuous conidiogenous cells, thickened and darkened. Conidia solitary, acicular, straight to curved, about 35–210 × 2–4 µm, 5–27-septate, hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate, hila thickened and darkened.

Holotype: India: Uttar Pradesh: without detailed locality, on Althernanthera sessilis, Amaranthaceae, Oct. 1992, S. Narayan (GPU 5023). Isotype: HCIO 41940.

Host range and distribution: On Alternanthera sessilis, Amaranthaceae, Asia (India).

Notes: This invalid species name pertains to the Cercospora apii s. lat. complex. The conidia were described to be “cylindrical” although obviously acicular conidia were depicted in the original drawing.

Cercospora crassoides Davis, Trans. Wisconsin Acad. Sci. 21: 298 (1924).

Synonym: Nimbya crassoides (Davis) E.G. Simmons, Mycotaxon 55: 146 (1995).

Literature: Chupp (1954: 33), Crous & Braun (2003: 143).

Illustration: Simmons (1995: 147, fig. 117).

Holotype: USA: Wisconsin: Lone Rock, on Froelichia floridana, 23 Jul. 1921, J. J. Davis (WIS). Isotypes: BPI 435265, CUP 39528.

Host range and distribution: On Froelichia floridana, Amaranthaceae, North America (USA, Florida, Oklahoma, Texas, Wisconsin).

Cercospora gomphrenae-globosae S. Narayan et al., in Rao et al., Sugarcane Pathology 1: Fungal Diseases: 69 (1999), nom. inval. (Arts 37.3 and 39.1).

Literature: Crous & Braun (2003: 2001), Kamal (2010: 47).

Illustration: Rao et al. (1999: 70, fig. 6).

Description: Leaf spots amphigenous, circular or subcicular, 1–10 mm diam, later to 30 mm diam, withish brown above, rusty brown below, with margin. Caespituli amphigenous, effuse. Mycelium internal; hyphae branched, septate, light olivaceous. Stromata well-developed, compact, subepidermal, about 10–30 µm diam, light olivaceous to brown. Conidiophores solitary or in small fascicles, 3–7, arising from stromata, erect, straight or almost so, unbranched, cylindrical, 1–4 times geniculate, about 45–155 × 3.5–5 µm, 2–7-septate, olivaceous to light brown, thin-walled, smooth; conidiogenous cells integrated, terminal and intercalary, conidiogenous loci conspicuous, thickened and darkened. Conidia solitary, subcylindrical to subacicular (somewhat attenuated towards the tip), straight to somewhat curved, about 13–205 × 2–3 µm, usually 7–18-septate, hyaline, thin-walled, smooth, apex subacute to obtuse, base truncate, thickened and darkened.

Syntypes (holotype not clearly indicated): India: Uttar Pradesh: Gorakhpur, on Gomphrena globosa, Nov. 1993/1994, S. Narayan (GPU 5051, 5061; HCIO 41968, 41978).

Host range and distribution: On Gomphrena globosa, Amaranthaceae, Asia (India, Uttar Pradesh).

Notes: This species is well-characterised by having subcylindrical conidia, but a validation is not made since type material was not available. The authors cited collections from two years, 1993 and 1994, but a holotype was not clearly designated. Furthermore, the name C. gomphrenaeglobosae is invalid since it was published without any Latin diagnosis or description.

Cercospora nothosaervae M.S. Patil, Botanique 6: 221 (1975).

Literature: Crous & Braun (2003: 292), Kamal (2010: 69).

Illustration: Patil (1975: 220, fig. 9).

Description: Leafspots circular 1–2 mm diam, centre depressed, whitish grey, margin red. Caespituli amphigenous. Mycelium internal. Stromata lacking. Conidiophores solitary or in small, loose groups, 2–3, erumpent, erect, straight to flexuous, non-geniculate, unbranched or occasionally branched, basal cells swollen, 20–70 × 5–5.5 µm, 1–2-septate, olivaceous-brown, paler towards the tip; conidiogenous loci conspicuous, prominent. Conidia solitary, broadly obclavate with attenuated, almost rostrate apex, straight to slightly curved, 20–70 × 5–6.5 µm, 1–5-septate, constricted at the septa, pale olivaceous-brown, apex subobtuse, base obconically truncate, with a prominent hilum, darkened and thickened.

Holotype: India: Maharashtra: Kolhapur, on Nothosaerva brachiata, Amaranthaceae, 17 Dec. 1973, M. S. Patil (HCIO 31692).

Host range and distribution: Only known from the type collections.

Notes: The general characters of this fungus are not cercosporoid. It seems to be a species of Alternaria, but it was not possible to confirm this assumption. Type material was not available.

Cercospora pretoriensis Chupp & Doidge, Bothalia 4: 890 (1948). (Fig. 73)

Fig. 73
figure73

Cercospora pretoriensis (CUP 40641, isotype). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm

Literature: Chupp (1954: 35), Crous & Braun (1996: 298), Crous & Braun (2003: 335).

Description: Leaf spots amphigenous, irregularly scattered, subcircular, 2–6 mm diam, sometimes confluent, on the upper surface centre pale brown to dingy grey, with reddish to red-brown border, on the lower surface paler. Caespituli hypophyllous, scattered to dense, punctiform, barely confluent, dark olivaceous. Mycelium internal. Stromata immersed, 15–60 µm diam, somewhat erumpent, olivaceous-brown, composed of swollen hyphal cells, angular-subglobose, about 3–9 µm diam. Conidiophores densely fasciculate, few to numerous, arising from stromata, erumpent, erect, subcylindrical, somewhat attenuated towards the tip, unbranched, straight to curved or usually distinctly, often strongly geniculate-sinuous, 10–50(− 100) × 3–6 µm, 0–3-septate, rarely pluriseptate, pale olivaceous-brown, paler towards the tip, upper part subhyaline or sometime pigmented throughout, thin-walled, smooth; conidiogenous cells integrated, terminal, sometimes conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci conspicuous or subconspicuous, 1.5–2 µm diam, visible in lateral view as convex to truncate tip or shoulder caused by sympodial proliferation (light microscopy), unthickened or only slightly thickened, not to slightly darkened, often somewhat refractive, in front view sometimes visible as minute circle with somewhat darker rim. Conidia solitary, narrowly cylindrical, occasionally subacicular, straight to slightly curved, 15–90 × 2–4.5 µm, 1- to pluriseptate, hyaline or subhyaline, thin-walled, smooth, apex obtuse or subobtuse, base truncate, subtruncate or slightly short obconically truncate, 1–2.5 µm wide, hila unthickened or only very slightly thickened, not darkened, at most somewhat refractive.

Holotype: South Africa: Transvaal: Pretoria, Arcadia, on Gomphrena globosa, 14 Apr. 1913, collector not indicated (PREM 6593). Isotype: CUP 40641. Paratypes: CUP40639, 40640; PREM 775, 26316, 32789.

Host range and distribution: On Gomphrena (celosioides, serrata [decumbens], globosa), Amaranthaceae, South Africa.

Notes: Cercospora pretoriensis is a cercosporoid species that cannot be unequivocally assigned to any of the currently recognised cercosporoid genera just based on morphology. The structure of the conidiogenous loci does not agree with scars of Cercospora s. str. and Passalora s. lat. Colourless conidia are also not in favour of the latter genus. The conidiogenous cells and conidiogenous loci are reminiscent of Cercosporella species, but the conidiophores are pigmented, often even throughout, which would be very unusual for Cercosporella. Moreover, relations to the complex around Paracercospora and Pseudocercosporella can also not be excluded with certainty. Phylogenetic data are required to elucidate the correct generic affinity of this species. For the interim, we prefer to maintain this species in Cercospora sensu latissimo.

Cercospora pseudachyranthis R.F. Castaneda & U. Braun, Cryptog. Bot. 1: 43 (1989). (Fig. 74)

Fig. 74
figure74

Cercosporella pseudachyranthis (HAL 1648 F, isotype). A. Conidiophore fascicle. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Synonym: Cercosporella pseudachyranthis (R.F. Castañeda & U. Braun) U. Braun, comb. nov. MycoBank MB814568

Basionym: Cercospora pseudachyranthis R.F. Castañeda & U. Braun, Cryptog. Bot. 1: 43 (1989).

Synonyms: Cercospora centrostachydis Chupp, Monograph of Cercospora: 32 (1954), nom. inval. (Art. 39.1) [type: Puerto Rico: Mayaguez, on Achyranthes aspera var. indica, 2 Mar. 1916, Whetzel & Olive 464].

Passalora pseudachyranthis (R.F. Castaneda & U. Braun) U. Braun, Nova Hedwigia 55: 221 (1992).

Misapplied name: Cercospora achyranthis sensu Solheim & Stevens (1931: 378).

Literature: Solheim & Stevens (1931: 378, as C. achyranthis), Chupp (1954: 32), Crous & Braun (2003: 337).

Illustration: Castañeda & Braun (1989: 44, pl. 1, fig. 1).

Description: Leaf spots amphigenous, subcircular to somewhat irregular, 0.5–4 mm diam, yellowish, ochraceous to brownish, margin indefinite or narrow and darker. Caespituli hypophyllous, punctiform-effuse, greyish white. Mycelium internal. Stromata well-developed, substomatal, subglobose, 20–40 µm diam, brownish, composed of swollen hyphal cells, 2–8 µm diam, subglobose-angular. Conidiophores in small to large fascicles (to 50 or even more), divergent to dense, arising from stromata, emerging through stomata, erect, straight to flexuous, curved, in the upper part slightly to mostly distinctly geniculate-sinuous, simple, rarely branched, (40−)50–200(− 220) × (2−)3–5(− 6) µm, septate, longer conidiophores pluriseptate, hyaline or subhyaline throughout to pale olivaceous below, occasionally slightly darker near the base, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–40 µm long, conidiogenous loci conspicuous, terminal or formed on lateral shoulders caused by sympodial proliferation, subtruncate to usually convex (light microscopy), slightly thickened, not darkened, but often somewhat refractive, 1.5–2.5 µm diam. Conidia solitary, obclavate-cylindrical, straight to slightly curved, 20–80 × 3.5–7 µm, 1–8-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, base obconically truncate to rounded, 1.5–2 µm wide, hila almost unthickened or only very slightly thickened, not darkened, but somewhat refractive.

Holotype: Cuba: San Miguel de los Baños, on Achyranthes aspera, 22 Dec. 1987, R.F. Castaneda (INIFAT C87/365). Isotype: HAL 1648 F. Paratypes: HAL 1761 F, INIFAT C87/375.

Host range and distribution: On Achyranthes (aspera var. aspera, aspera var. indica [indica, Centrostachys indica], bidentata, Achryranthes sp.), Amaranthaceae, ?Asia (Pakistan), West Indies (Cuba, Puerto Rico, Virgin Islands).

Notes: The morphological characters of this species have previously been misinterpreted. Due to colourless conidia and conspicuous conidiogenous loci, this species was originally assigned to Cercospora (Castañeda & Braun 1989), but later reallocated to Passalora (Braun 1992). However, the structure of the conidiogenous loci does not coincide with the concepts of the two genera, and colourless conidia are not consistent with the current concept of Passalora. Type material of this species has been re-examined and revealed that the conidia and the structure of conidiogenous cells and conidiogenous loci are in agreement with Cercosporella (Braun 1995). The conidiophores are not quite colourless, above all in the lower half, but this kind of pigmentation is known in some tropical-subtropical species of Cercosporella (Braun 1995).

Chupp (1954) introduced the new species name Cercospora centrostachydis with reference to Solheim & Stevens (1931: 378, as C. achyranthis), but failed to add a Latin description or diagnosis which was necessary for a valid publication in 1954. The type material cited in Chupp (1954), not agreeing with the collections cited in Solheim & Stevens (1931), was not traced in Chupp’s herbarium in CUP. A record of C. centrostachydis from Pakistan (Ahmad et al. 1997) is unclear and unproven.

Cercospora pupaliae Patwardhan & A.K. Pande, Sydowia 23: 98 “1969” (1970); as “pupalae”.

Literature: Crous & Braun (2003: 344), Kamal (2010: 79).

Illustration: Patwardhan & Pande (1970: 99, fig. 6).

Description: Leaf spots circular to irregular, grey to dirty brown, margin brown to black, mostly epiphyllous. Stromata 10–30 µm diam, dark brown. Conidiophores in fascicles of 4–10, arising from stromata, divergent, unbranched, 18.5–96.6 × 3.7 µm, darker brown below, paler or subhyaline above. Conidia solitary, obclavate, straight, 51.8–70 × 3.7 µm, septate, hyaline to olivaceous.

Holotype: India: Maharashtra: Pune, on Pupalia orbiculata, Amaranthaceae, 23 Aug. 1966, G. P. Patwardhan (AMH 294).

Host range and distribution: Only known from the type collection.

Notes: Type material of this species was not available. The original description is meagre. Kamal (2010) alluded that he had examined type material of C. pupaliae. He emphasized that that this species is distinctly different from C. apii, but he did not provide any description or illustration and did not specify these differences, above all he did not refer to the original description of conidia (hyaline to olivaceous), although olivaceous conidia could be an indication for Passalora s. lat., i.e. based on the original description and illustration, this species might belong to the latter genus. Furthermore, the conidia were described to be “acicular” but the illustration shows them to be obclavate. Hence, the generic affinity of C. pupaliae is not quite clear and this species is in urgent need of revision.

Cercospora sessilis Pavgi & U.P. Singh, Mycopathol. Mycol. Appl. 23: 190 (1964), nom. illeg. (Art. 53.1), non C. sessilis Sorokin, 1892.

Literature: Crous & Braun (2003: 375), Kamal (2010: 85).

Illustration: Pavgh & Singh (1964:192, figs 1012).

Description (based on Pavgh & Singh 1964 and examination of type material): Leaf spots amphigenous, greenish brown, on the upper surface becoming chlorotic, 4–8 mm diam, confluent. Stromata substomatal, 10–25 µm diam, pale olivaceous-brown. Conidiophores in small to moderately large fascicles, divergent to moderately dense, arising from stromata, erect, straight, subcylindrical to somewhat geniculate-sinuous, unbranched, 25–90 × 3–4 µm, continuous to septate, light brown, thin-walled, smooth; conidiogenous cells integrated, terminal. Conidia solitary, obclavate, 30–120 × 1.5–3 µm, 3–11-septate, hyaline or subhyaline, thin-walled, smooth, apex subacute, base short obconically truncate.

Syntypes: India: Uttar Pradesh: Varanasi, on Alternanthera sessilis, Amaranthaceae, 22 Oct. 1961, U. P. Singh (HCIO, K(M) IMI 113095).

Notes: Status and generic affinity unclear, type material deposited at K(M) has been examined, but it was too meagre for a final conclusion. According to the original publication, syntype material has been deposited in HCIO, but an accession number was not cited and the material concerned was not available. Some conidiophores were found, all without any conspicuous conidiogenous loci, and a single conidium which seemed to have a slightly thickened and darkened hilum.

Passalora

Key to Passalora species on Amaranthaceae
  1. 1

    Conidia formed singly, obclavate-cylindrical, 30–85 × 3.5–6 µm, (1−)3–7-septate, pale olivaceous; mycelium internal; conidiophores fasciculate, arising from well-developed stromata, 25–75 µm diam; on Pfaffia sericea ……….. P. pfaffiae

    Conidia at least partly catenate and/or superficial mycelium with solitary conidiophores formed and stromata lacking ……….. 2

  2. 2(1)

    Superficial hyphae with solitary conidiophores formed; stromata lacking (mycovellosiella-like species) ……….. 3

    Superficial hyphae and solitary conidiophores lacking; conidiophores fasciculate; stromata developed (phaeoramularia-like species) ……….. 4

  3. 3(2)

    Conidiophores relatively long, 10–200 × 3–7 µm, longer conidiophores pluriseptate; on Cyathula achyranthoides ……….. P. cyathulae

    Conidiophores shorter, 5–30 × 3–5 µm, 0–1-septate; on Iresine spp ……….. P. gonoclada

  4. 4(2)

    Conidia usually cylindrical or subcylindrical, hyaline or subhyaline, 20–60 × 2–5 µm, 0–5-septate; conidiophores narrow, 1.5–5 µm wide; on Iresine diffusa ……….. P. gilbertii

    Conidial shape variable, cylindrical, fusiform to obclacate, olivaceous to olivaceous-brown, 3–8 µm wide; conidiophores broader, 2–8 µm ……….. 5

  5. 5(4)

    Conidiophores occasionally branched; on Iresine diffusa ……….. P. iresines

    Conidiophores unbranched; on Gomphrena and Pfaffia spp. ……….. P. gomphrenicola

Tabular key to Passalora species on Amaranthaceae according to host genera
  • Cyathula

    A single species ……….. P. cyathulae

  • Gomphrena

    A single species ……….. P. gomphrenicola

  • Iresine

    1. 1

      Stromata lacking; conidiophores solitary, arising from superficial hyphae, 5–30 µm long, 0–1-septate ……….. P. gonoclada

      Stromata developed; superficial hyphae and solitary conidiophores absent; conidiophores fasciculate, 20–60(− 110) µm long, 0–5-septate ……….. 2

    2. 2(1)

      Conidia usually cylindrical or subcylindrical, hyaline or subhyaline, 20–60 × 3.5–6 µm; conidiophores narrow, 1.5–5 µm wide, unbranched ……….. P. gilbertii

      Conidial shape variable, cylindrical, fusiform to obclacate, olivaceous to olivaceous-brown, 3–8 µm wide; conidiophores broader, 4–6(− 8) µm, occasionally branched ……….. P. iresines

  • Pfaffia

    1. 1

      Conidia in chains, ellipsoid, ovoid, broadly fusiform, subcylindrical, (10−)15–60 × 3–8 µm, (0−)1–4(− 5)-septate ……….. P. gomphrenicola

      Conidia solitary, obclavate-cylindrical, 30–85 × 3.5–6 µm, (1−)3–7-septate ……….. P. pfaffiae

Passalora species on Amaranthaceae

Passalora cyathulae (F. Stevens & Solheim) U. Braun & Crous, Mycosphaerella and Anam.: 148 (2003) (Fig. 75)

Fig. 75
figure75

Passalora cyathulae (ILL 11981, holotype). A. Superficial hyphae. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophore. D. Conidia. Bar = 10 µm.

Basionym: Ragnhildiana cyathulae F. Stevens & Solheim, Mycologia 23: 403 (1931)

Literature: Chupp (1954: 33).

Illustration: Solheim & Stevens (1931: 403, fig. 12).

Description: Leaf spots lacking or indistinct, yellowish discolorations or small circular, subcircular to somewhat irregular spots, 0.5–2 mm diam, brown above, olivaceous-brown below, margin indefinite. Caespituli hypophyllous, effuse, dark olivaceous. Mycelium internal and external; superficial hyphae emerging through stomata, branched, septate, 2–5 µm wide, subhyaline to pale brown, thin-walled, smooth. Stromata lacking. Conidiophores solitary, arising from superficial hyphae, lateral or terminal, erect to decumbent (differentiation between solitary conidiophores arising from superficial hyphae and long decumbent conidiophores difficult), straight, subcylindrical to geniculate-sinuous, unbranched to branched, length variable, 10–200 × 3–7 µm, aseptate to pluriseptate throughout, subhyaline to pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, sometimes conidiophores reduced to conidiogenous cells, 10–30 µm long, conidiogenous loci conspicuous, thickened and darkened, (1.5−)2(− 2.5) µm diam. Conidia in simple or sometimes branched chains, straight to somewhat curved, ellipsoid to cylindrical, 20–65 × 3–7.5 µm, 0–4-septate, subhyaline to pale brownish, thin-walled, smooth, ends short obconically truncate to rounded, about 2 µm wide, hila somewhat thickened and darkened.

Holotype: Guyana: Coverden, on Cyathula achyranthoides, Amaranthaceae, 4 Aug. 1922, F. L. Stevens 743 (ILL 11981).

Host range and distribution: Only known from the type collection.

Notes: This species is a typical mycovellosiella-like Passalora species with superficial hyphae with solitary conidiophores, thickened and darkened conidiogenous loci and conidial hila, and conidia formed in chains. Chupp (1954) confused Ragnhildiana cyathulae and the Indian Cercospora cyathulae described by Sydow (1937). Furthermore, he cited “Cercospora cyathulae (F. Stevens & Solheim) Syd.” which is, however, incorrect. Sydow (in Sydow et al. 1937) described a new Indian species but did not introduced a new species based on Ragnhildiana cyatheae. Moreover, Chupp’s (1954) description of C. cyathulae was based on characters of both species, although the two species are readily distinguishable by obvious differences in the conidial length and septation.

Passalora gilbertii (Speg.) U. Braun, Schlechtendalia 5: 62 (2000). (Fig. 76)

Fig. 76
figure76

Passalora gilbertii (Syd., Fungi Exot. Exs. 1047, HBG, neotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora gilbertii Speg., Anales Soc. Ci. Argent. 10(1): 32 (1880).

Synonym: Phaeoramularia gilbertii (Speg.) U. Braun, Schlechtendalia 2: 11 (1999).

Literature: Saccardo (1886: 457), Chupp (1954: 33), Crous & Braun (2003: 197).

Illustration: Braun (1999: 12, fig. 14).

Exsiccatae: Syd., Fungi Exot. Exs. 1047.

Description: Leaf spots amphigenous, subcircular to irregular, 1–10 mm diam, oblong patches to 20 mm in length, brown to dingy grey, zonate, sometimes with narrow brown margin. Caespituli amphigenous, punctiform, dark brown to blackish. Mycelium internal. Stromata well-developed, 20–60 µm diam, subglobose, immersed, brown. Conidiophores numerous, in loose to very dense fascicles, arising from stromata, erumpent, erect, filiform, flexuous, somewhat geniculate-sinuous, unbranched, 20–60 × 1.5–5 µm, aseptate to pluriseptate throughout, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–30 µm long, conidiogenous loci subconspicuous, minute, 0.75–1.5 µm diam, slightly thickened and darkened-refractive. Conidia solitary or catenate, in short chains, subcylindrical, occasionally narrowly obclavate to fusiform, 20–60 × 2–5 µm, (0−)1–4-septate, hyaline or subhyaline, thin-walled, smooth, apex obtuse, subacute or truncate in catenate conidia, base short to long obconically truncate, about 1 µm wide, hila barely thickened, colourless to slightly darkened-refractive.

[Holotype: Uruguay: Montevideo, on Iresine diffusa, 1876, G. Gilbert 908 (not preserved).] Neotype (designated by Crous & Braun 2003): Ecuador: Quito, Pichincha Mt., 30 Sep. 1937, H. Sydow [Syd., Fungi Exot. Exs. 1047] (HBG). Isoneotypes: Syd., Fungi Exot. Exs. 1047 (e.g. BPI 436666).

Host range and distribution: On Iresine diffusa [celosia, celosioides, paniculata], Amaranthaceae, Africa (São Tomé e Príncipe), North America (Mexico), South America (Colombia, Ecuador, Uruguay), West Indies (Puerto Rico, Virgin Islands).

Note: The orthographic variant “gibertii” sometimes used for this species is incorrect. The epithet was derived from the name of the collector, G. Gilbert. The phylogenetic affinity of Cercospora gilbertii, characterised by an unusual combination of morphological traits, is quite uncertain. Colourless conidia are in favour of Cercospora in its current circumscription, but catenate conidia and the structure of the conidiogenous loci argue against it. Molecular analyses are necessary to elucidate the true generic affinity of this species. For the interim we prefer to maintain this species in Passlora s. lat.

Passalora gomphrenicola (Speg.) U. Braun, Schlechtendalia 5: 64 (2000). (Fig. 77)

Fig. 77
figure77

Passalora gomphrenicola (LPS 914, holotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from superficial hypha. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Cercospora gomphrenicola Speg., Anales Soc. Ci. Argent. 13: 29 (1882).

Synonym: Phaeoramularia gomphrenicola (Speg.) Munt.-Cvetk., Lilloa 30: 209 (1960).

Literature: Saccardo (1886: 457), Chupp (1954: 34), Ellis (1971: 308), Deighton (1979: 27), Pons & Sutton (1988: 31), Crous & Braun (2003: 201–202).

Illustrations: Muntañola (1960: 210–211, figs 1617), Ellis (1971: 307, fig. 213), Deighton (1979: 27, fig. 13).

Exsiccatae: Speg., Hongos Sud-Amer. Dec. Mycol. Argent. 45.

Description: Leaf spots lacking or only visible as yellowish discolorations, turning dark olivaceous by abundant formation of caespituli, patches to 12 mm diam. Caespituli hypophyllous, effuse, olivaceous, forming small patches or confluent. Mycelium internal, hyphae colourless, branched, septate, 2–4 µm wide. Stromata absent or small, about 10–70 µm diam, substomatal, pale to brown, composed of swollen hyphal cells, circular to somewhat angular-irregular in outline, 2–6 µm diam. Conidiophores in small to large, loose to dense fascicles, larger fascicles composed of 50 or even more conidiophores, arising from internal hyphae or from stromata, through stomata or erumpent, erect to decumbent, straight, subcylindrical-conical to sinuous or somewhat geniculate-sinuous, unbranched or branched, decumbent branched conidiophores reminiscent of and confusable with superficial hyphae giving rise to solitary conidiophores, 20–70(− 90) × 2–7 µm, often irregular in width, 1–5-septate, often somewhat constricted at the septa, pale olivaceous to olivaceous-brown, darker in mass, thin-walled, smooth; conidiogenous cells integrated, terminal, intercalary and occasionally pleurogenous, 5–30 µm long, proliferation sympodial, occasionally percurrent, conidiogenous loci conspicuous, sometimes subdenticulate, thickened and darkened, 1.5–2.5 µm wide. Conidia in simple or branched chains, ellipsoid, ovoid, broadly fusiform, subcylindrical, straight to slightly curved, (10−)15–60 × 3–8 µm, (0−)1–4(− 5)-septate, occasionally somewhat constricted at the septa, pale olivaceous to olivaceous-brown, thin-walled, smooth, occasionally faintly rough, apex rounded to truncate in catenate conidia, base short obconically truncate, (1.5−) 2(− 2.5) µm wide, hila somewhat thickened and darkened.

Holotype: Argentina: Buenos Aires, Palermo, on Pfaffia glomerata, Feb. 1881, C. Spegazzini (LPS 914). Isotypes: Speg., Hongos Sud-Amer. Dec. Mycol. Argent. 45, e.g. BPI 436740, 722393; K(M) IMI 7706 (slide), MICH 15302.

Host range and distribution: On Gomphrena globosa, Pfaffia (glomerata [glauca, stenophylla; Gomphrena glauca], iresinoides), Amaranthaceae, Africa (South Africa, Transvaal), South America (Argentina, Brazil, Venezuela).

Passalora gonatoclada (Syd.) U. Braun & Crous, Mycosphaerella and Anam.: 202 (2003). (Fig. 78)

Fig. 78
figure78

Passalora gonatoclada (Syd., Fungi Exot. Exs. 930, CUP, isolectotype). A. Solitary conidiophores arising from superficial hyphae and through a stoma. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora gonatoclada Syd., Ann. Mycol. 23: 425 (1925).

Synonyms: Ragnhildiana gonatoclada (Syd.) F. Stevens & Solheim, Mycologia 23: 403 (1931).

Mycovellosiella gonatoclada (Syd.) Munt.-Cvetk., Lilloa 30: 106 & 208 (1960), nom. inval. (Art. 41.5).

Mycovellosiella gonatoclada (Syd.) Deighton, Mycol. Pap. 137: 69 (1974).

Literature: Chupp (1954: 35), Deighton (1974: 69), Crous & Braun (2003: 202).

Illustration: Muntañola (1960: 208, fig. 15 A).

Description: Leaf spots indistinct, formed as yellowish discolorations on the upper leaf surface. Colonies hypophyllous, effuse, brownish. Mycelium internal and external; superficial hyphae solitary, occasionaly intertwined, branched, septate, subhyaline to olivaceous-brown, 2–8 µm wide (sterile hyphae paler and narrow, fertile hyphae with conidiophores broader and darker), thin-walled, smooth. Stromata lacking. Conidiophores solitary, arising from superficial hyphae, lateral or terminal, erect to decumbent, 5–30 × 3–10 µm, 0–1-septate (decumbent threats with terminal conidiophores may be much longer and pluriseptate), sometimes with intercalary cells giving rise to minute peg-like protuberances with a single terminal scar, only about 2–5 µm long and wide, subhyaline to pale olivaceous or olivaceous-brown, thin-walled, smooth; conidiophores mostly reduced to conidiogenous cells, conidiogenous loci conspicuous, somewhat thickened and darkened, 1.5–2 µm diam. Conidia catenate, in simple or branched chains, ellipsoid-ovoid to cylindrical, 12–60 × 3.5–8.5 µm, (0−)1–7-septate, subhyaline to pale olivaceous-brown or brownish, thin-walled, smooth, apex rounded in terminal primary conidia or conically truncate in secondary (catenate) conidia, subdenticulate when in branched chains, base short obconically truncate, 1.5–2.5 µm wide, hila thickened and darkened.

Lectotype (designated here, MycoBank, MBT202803): Costa Rica: San José, La Caja, on Iresine calcea, 7 Jan. 1925, H. Sydow [Syd., Fungi Exot. Exs. 930] (S-F57194). Isolectotypes: Syd., Fungi Exot. Exs. 930, e.g., BPI 436742, CUP, HBG, K(M) IMI 7704, MICH 15303, NY 937030. Former syntypes (type locality but from 5 Jan. 1925 [H. Sydow, Fungi itin. Costaricensi Coll. 12]): E 417850, ILL 10671.

Host range and distribution: On Iresine (calea, diffusa [paniculata]), Amaranthaceae, Central America (Costa Rica), West Indies (Puerto Rico, Virgin Islands).

Passalora iresines (Munt.-Cvetk.) U. Braun & Crous, Mycosphaerella and Anam.: 456 (2003) (Fig. 79)

Fig. 79
figure79

Passalora iresines (based on Muntañola 1960: 217, fig. 20, 219, fig. 21). A. Conidiophore fascicles. B. Conidiophore tips. C. Conidia. Bar = 10 µm.

Basionym: Phaeoramularia iresines Munt.-Cvetk., Lilloa 30: 216 (1960).

Illustrations: Muntañola (1960: 217, fig. 20, 219, fig. 21).

Description: Leaf spots formed as epiphyllous chlorotic discolorations, scattered to confluent, circular, elliptical, diffuse, violaceous, finally ochraceous to brown. Caespituli hypophyllious, velutinous, brownish to brown-olivaceous, patches subcircular or sometimes vein-limited. Mycelium internal. Stromata moderately large, substomatal, olivaceous. Conidiophores in divergent to dense fascicles, arising from stromata, through stomata, erect, Aexuous, geniculate-sinuous, simple or occasionally branched, 40–60 (− 110) × 4–6(− 8) µm, branchlets 15–20(− 50) µm long, base sometimes swollen, aseptate to sparingly septate, often somewhat constricted at septa, olivaceous, paler towards the tip, subhyaline, thin-walled, smooth; conidiogenous cells integrated, terminal or intercalary, sometimes lateral, or conidiophores reduced to conidiogenous cells, conidiogenous loci conspicuous, thickened and darkened. Conidia catenate, in simple or branched chains, variable in shape and size, cylindrical, fusiform, obclavate, straight to curved, rarely sigmoid, 20–60 × 3.5–6 µm, 0–3-septate, olivaceous, thin-walled, smooth, base obconically truncate, with a single somewhat thickened and darkened hilum, apex rounded or with 1–3 hila.

Holotype: Argentina: Tucumán: El Cerro San Javier, on Iresine diffusa, 1 Aug. 1959, M. Muntañola (not traced).

Host range and distribution: On Iresine diffusa [polymorpha], Amaranthaceae, South America (Argentina).

Note: Muntañola (1960) designated a collection in her private herbarium (no. 600) as type. Muntañola-Cvetković died in 2011. The fate of her herbarium could not yet be clarified. After her time in South America, she worked in Serbia (University of Belgrade, Faculty of Biology), and after her retirement she moved to Portugal (University of Barcelona, Faculty of Biology), but Muntañola’s herbarium is not preserved in Belgrad (J. Vukojevic, pers. comm.). A part of her herbarium, mainly fungi collected in Portugal, are preserved in BCN, but South American collections are not included (according to the curator of BCN, Barcelona).

Passalora pfaffiae (Chupp) U. Braun & Crous, Mycosphaerella and Anam.: 318 (2003). (Fig. 80)

Fig. 80
figure80

Passalora pfaffiae (CUP 40533, holotype). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora pfaffiae Chupp, Monograph Cercospora: 35 (1954).

Literature: Chupp (1954: 35).

Description: Leaf spots amphigenous, circular to somerwhat angular-irregular, 1–6 mm diam, pale greyish brown to dingy grey, with narrow marginal line, somewhat raised. Caespituli amphigenous, punctiform, scattered, dark brown to blackish. Mycelium internal. Stromata substomatal to immersed, subglobose to oblong, 25–75 µm diam, dark brown, composed of subglobose cells, 2.5–5 µm diam. Conidiophores in moderately large fascicles, divergent to mostly dense, arising from stromata, erect, subcylindrical, sinuous or slightly geniculate, unbranched, 10–70 × 3.5–6 µm, 0–3-septate, pale olivaceous to brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, 10–30 µm long, proliferation sympodial, occasionally percurrent, with fine annellations, conidiogenous loci conspicuous, small, 1–1.5 µm diam, somewhat thickened and darkened. Conidia solitary, obclavate-cylindrical, straight to somewhat curved, 30–85 × 3.5–6 µm, (1−)3–7-septate, pale olivaceous, thin-walled, smooth, apex obtuse, base short obconically truncate, 1.5–2 µm wide, hila somewhat thickened and darkened.

Holotype: Brazil: Rio Grande do Sul: Taquari, Parque Apicola, on Pfaffia sericea, Amaranthaceae, 31 Dec. 1946, J. P. da Costa Neto 2224 (CUP 40533).

Host range and distribution: Only known from the type collection.

Note: Based on a combination of conspicuous conidiogenous loci, slightly thickened and darkened, 1–1.5 µm wide, and obclavate-cylindrical, pale olivaceous conidia, Crous & Braun (2003) reallocated this species to Passalora.

Pseudocercospora

Key to Pseudocercospora species on Amaranthaceae
  1. 1

    Superficial hyphae with solitary conidiophores developed ……….. 2

    Superficial hyphae with solitary conidiophores lacking; conidiophores consistently fasciculate ……….. 4

  2. 2(1)

    Conidia broad, 4–8 µm; on Cyathula tomentosa ……….. P. cyathulae

    Conidia narrower, 2–4 µm ……….. 3

  3. 3(2)

    Conidia obclavate-cylindrical, pale olivaceous-brown; on Alternanthra spp ……….. P. alternantherae

    Conidia cylindrical, subacicular to obclavate-cylindrical, subhyaline to pale olivaceous; on Celosia spp ……….. P. celosiarum

  4. 4(1)

    Conidiophores 10–100 µm long; conidia narrowly filiform-subcylindrical, subacicular or narrowly obclavate-cylindrical, (40−)60–130(− 150) × 1.5–3.5 µm, 2–12-septate, hila 1–1.5 µm wide; on Gomphrena spp. ……….. P. globosae

    Conidiophores much shorter, 10–35 µm long; conidia obclavate-cylindrical, 10–90 µm long, 0–10-septate or, if conidiophores longer, conidia 2.5–6 µm wide; on other hosts ……….. 5

  5. 5(4)

    Conidiophores 10–70 × 3–5.5 µm, 0–5-septate; conidia 25–135 × 2.5–6 µm, 3–16-septate, pale olivaceous-brown; on Alternanthera spp. ……….. P. alternantherae-nodosae

    Conidiophores shorter, 5–35 µm long, only 0–1(− 2)-septate; conidia shorter and narrower, 15–90 × 2–4 µm or subhyaline if broader; on other hosts ……….. 6

  6. 6(5)

    Stromata lacking; conidiophores in small fascicles; on Gomphrena spp ……….. P. gomphrenae

    Stromata developed, 10–100 µm diam; conidiophores mostly in larger fascicles arising from stromata ……….. 7

  7. 7(6)

    Stromata large, 10–100 µm diam; conidia (2−)3–5(− 6) µm wide, hila 1.5–3 µm wide, subhyaline; on Gomphrena pulchella ……….. P. gomphrenae-pulchellae

    Stromata smaller, 10–65 µm; conidia 2–3.5 µm wide, hila 1–2 µm wide, subhyaline to pale olivaceous-brown; on other hosts ……….. 8

  8. 8(7)

    Conidia mostly narrowly cylindrical with truncate base, pale olivaceous-brown; on Amaranthus spp ……….. P. amaranthicola

    Conidia obclavate-cylindrical, base short obconically truncate, subhyaline to pale olivaceous; on other hosts ……….. 9

  9. 9(8)

    Forming distinct leaf spots; caespituli hypophyllous, punctiform; stromata substomatal; on Chamissoa altissima ……….. P. chamissoana

    Leaf spots lacking or only with indistinct discolorations; caespituli epiphyllous, forming sooty patches; stromata immersed; on Froelichia sp. ……….. P. froelichiae

Tabular key to Pseudocercospora species on Amaranthaceae according to host genera
  • Alternanthera

    1. 1

      Superficial hyphae with solitary conidiophores developed ……….. P. alternantherae

  • Superficial hyphae and solitary conidiophores absent; conidiophores only in fascicles ……….. P. alternantherae-nodosae

  • Amaranthus

    A single species ……….. P. amaranthicola

  • Celosia

    A single species ……….. P. celosiarum

  • Chamissoa

    A single species ……….. P. chamissoana

  • Cyathula

    A single species ……….. P. cyathulae

  • Froelichia

    A single species ……….. P. froelichiae

  • Gomphrena

    1. 1

      Stromata lacking, conidiophores in small fascicles ……….. P. gomphrenae

      Stromata developed, 10–100 µm diam; conidiophores in small to large fascicles, arising from stromata ……….. 2

    2. 2(1)

      Stromata 10–100 µm diam; conidiophores 10–30 × 2.5–6 µm; conidia obclavate-cylindrical, (2−)3–5(− 6) µm wide, subhyaline; on Gomphrena pulchella ……….. P. gomphrenae-pulchellae

      Stromata 10–40 µm diam; conidiophores 10–100 × 1.5–5.5 µm; conidia filiform-subcylindrical, subacicular to narrowly obclavate-cylindrical, narrower, 1.5–3.5 µm, subhyaline to pale olivaceous-brown; on Gomphrena globosa ……….. P. globosae

Pseudocercospora species on Amaranthaceae

Pseudocercospora alternatherae J.M. Yen et al., Mycotaxon 16: 39 (1982). (Fig. 81)

Fig. 81
figure81

Pseudocercospora alternantherae (based on Yen et al. 1982: 40, fig. 3). A. Conidiophore fascicles. B. Solitary conidiophores arising from superficial hyphae. C. Conidia. Bar = 10 µm.

Literature: Kamal (2010: 147).

Illustration: Yen et al. (1982: 40, fig. 3).

Description: Leaf spots amphigenous, scattered, oval to fusiform, 3–12 × 2–4 mm, often confluent, greyish brown, margin indistinct. Caespituli amphigenous, mostly hypophyllous, not very conspicuous. Mycelium internal and external, superficial; hyphae branched, septate, olivaceous-brown, thin-walled, smooth, 2–5 µm wide, internal hyphae 2–5 µm wide, external hyphae 2–3.5 µm wide. Stromata globose, substomatal, 20–40 µm diam, brown to dark brown. Conidiophores in small to well-developed fascicles, about 5–60, divergent to dense, arising from stromata, through stomata, and solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical to sinuous, slightly geniculate, unbranched, 15–55 × 3–4.5 µm, 0–3-septate, pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, obclavate-cylindrical, straight to slightly curved, 30–90 × 2.5–4 µm, 3–10-septate, pale olivaceous-brown, thin-walled, smooth, apex obtuse, base short obconically truncate, hila unthickened, not darkened.

Holotype: India: West Bengal: Purulea, on Alternanthera sp., Amaranthaceae, 7 Jan. 1981, B. K. Das Pcc4483 [Yen 10583] (not traced).

Host range and distribution: Only known from the type collection.

Notes: Yen et al. (1982) cited “LAM, Yen #10583” as holotype. The mycological LAM collections are now housed at UC, but the type material concerned could currently not be traced. It is possible that this material is still among numerous unincorporated Yen collections in UC (unnamed, only provided with a collection number).

Pseudocercospora alternantherae-nodiflorae (Sawada) Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 2: 135 (1987). (Fig. 82)

Fig. 82
figure82

Pseudocercospora alternantherae-nodiflorae (based on Hsieh & Goh (1990: 18, fig. 7). A. Conidiophore fascicle. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora alternantherae-nodiflorae Sawada, Rep. Gov. Agric. Res. Inst. Taiwan 35: 106 (1928).

Synonyms: Cercospora alternanthericola Pavgi & U.P. Singh, Mycopathol. Mycol. Appl. 27: 93 (1965) [holotype: India:

Uttar Pradesh: Varanasi, on Alternanthera sessilis, 17 Aug. 1963, U. P. Singh (MSP 273)].

Pseudocercospora alternanthericola (Pavgi & U.P. Singh) Deighton, Mycol. Pap. 140: 139 (1976).

Literature: Chupp (1954: 31), Hsieh & Goh (1990: 16), Guo & Hsieh (1995: 8), Guo et al. (1998: 19), Crous & Braun (2003: 53), Kamal (2010: 147).

Illustrations: Pavgi & Singh (1965: 94, pl. 1, figs 1618), Hsieh & Goh (1990: 18, fig. 7), Guo & Hsieh (1995: 9, fig. 8), Guo et al. (1998: 19, fig. 8).

Description: Leaf spots at first indistinct or visible as small brown spots, about 2–4 mm diam, later lower leaf surface gradually turning brown, upper leaf surface also discoloured, leaves finally disfigured, faded, necrotic. Caespituli amphigenous, forming effuse, dark olivaceous patches, finally covering the whole leaf blade. Mycelium internal. Stromata small, substomatal, 10–20 µm diam, brown. Conidiophores in small to well-developed fascicles, 6–25, loose to moderately dense, arising from stromata, through stomata, erect, straight, subcylindrical to curved or distinctly geniculate-sinuous, unbranched, 10–70 × 3–5.5 µm, 0–5-septate, uniformly pale olivaceous-brown to brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous to visible as truncate or conically truncate tips or shoulders formed by sympodial proliferation, unthickened, not darkened. Conidia solitary obclavate-subcylindrical, straight to curved, 25–135 × 2.5–6 µm, 3–15-septate, pale olivaceous, thin-walled, smooth, apex obtuse, rounded to subacute, base obconically truncate, 2–2.5 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202804): Taiwan: Taipei, on Alternanthera sessilis, 9 May 1924, K. Sawada (NTU-PPE, hb. Sawada). Isolectotypes: HMAS 05136, TNS-F-220608.

Host range and distribution: On Alternanthera sessilis [nodiflora], Alternanthera sp., Amaranthaceae, Asia (China; India, Uttar Pradesh; Taiwan).

Notes: Three duplicates of an additional syntype collection from Taiwan have been examined [Taipei, on Alternanthera sessilis, 5 Aug. 1907, K. Kawakami (BPI 432466, 432467; K(M) IMI 31945). Cercospora alternathericola, described from India on Alternanthera sessilis is barely distinct from Pseudocercospora alternantherae-nodiflorae and is therefore reduced to synonymy.

Pseudocercospora amaranthicola (J.M. Yen) J.M. Yen, Bull. Trimestriel Soc. Mycol. France 94: 385 (1978). (Fig. 83)

Fig. 83
figure83

Pseudocercospora amaranthicola (based on Yen 1977: 146, fig. 1 A–C). A. Conidiophore fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora amaranthicola J.M. Yen, Bull. Trimestriel Soc. Mycol. France 93: 145 (1977).

Synonym: Cercoseptoria amaranthicola (J.M. Yen) J.M. Yen, Bull. Trimestriel Soc. Mycol. France 97: 91 (1981); “as amaranticola”.

Literature: Hsieh & Goh (1990: 17), Guo & Hsieh (1995: 357), Guo et al. (1998: 376), Crous & Braun (2003: 55).

Illustrations: Yen (1977: 146, fig. 1A-C), Guo et al. (1998: 377, fig. 307).

Description: Leaf spots circular or subcircular, 0.5–4 mm diam, pale brown, margin brown. Caespituli epiphyllous, rarely amphigenous, punctiform, dark brown to blackish. Mycelium internal. Stromata immersed, globose or subglobose, 25–65 µm diam, dark brown. Conidiophores in well-developed, dense fascicles, arising from stromata, erumpent, erect, straight, flexuous, subcylindrical to geniculate-sinuous, unbranched or occasionally branched, 8–30 × 2.5–3.5 µm, 0–3-septate, pale olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous or visible as truncate tips or lateral shoulders caused by sympodial proliferation. Conidia solitary, narrowly cylindrical or only slightly obclavate-cylindrical, straight to curved, 15–80 × 2–3 µm, 3–10-septate, pale olivaceous-brown, apex obtuse or subobtuse, base truncate to somewhat obconically truncate, hila neither thickened nor darkened.

Holotype: Taiwan: Peikuoshan, Yuanlin, Changhua Hsien, on Amaranthus tricolor [mangostanus], Amaranthaceae, 30 Oct. 1971, J. M. Yen 71282.

Host range and distribution: Only known from the type collection.

Notes: Type material of this species could not be traced, neither in PC nor UC. It is possible that the material concerned is still among the numerous untreated Yen collection in the latter herbarium.

Pseudocercospora celosiarum (A.K. Kar & M. Mandal) Deighton, Mycol. Pap. 140: 141 (1976). (Fig. 84)

Fig. 84
figure84

Pseudocercospora celosiarum (K(M) IMI 135869, holotype). A. Conidiophore fascicles. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophores D. Conidia. Bar = 10 µm.

Basionym: Cercospora celosiarum A.K. Kar & M. Mandal, Trans. Brit. Mycol. Soc. 54: 423 (1970).

Literature: Katsuki (1965: 9, as “C. celosiae”), Guo & Hsieh (1995: 9), Guo et al. (1998: 20), Nakashima et al. (2002: 98), Crous & Braun (2003: 114), Kamal (2010: 162).

Illustrations: Kar & Mandal (1970: 423, fig. 1), Guo & Hsieh (1995: 10, fig. 9), Guo et al. (1998: 20, fig. 9).

Description: Leaf spots amphigenous, subcircular to angular-irregular or diffuse, size variable, 1.5–30 mm diam, brown, dull brown to dark greyish brown, margin narrow, darker brown to reddish brown. Caespituli amphigenous, punctiform, dark brown or more greyish by abundant conidiation. Mycelium internal and external; superficial hyphae, if present, emerging through stomata, branched, septate, 2–3 µm diam, subhyaline, thin-walled, smooth. Stromata almost lacking to well-developed, substomatal, subglobose, sometimes oblong, 5–60 µm diam, pale olivaceous to brown. Conidiophores in small to large fascicles, divergent to very dense, arising from stromata, through stomata or solitary, arising from superficial hyphae, lateral, erect, straight, subcylindrical-conical, not to strongly geniculate-sinuous, unbranched, 5–75 × 2–4 µm, 0–5-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous, neither thickened nor darkened. Conidia solitary, cylindrical, subacicular, long acicular to obclavate-cylindrical, 10–105 × 2–4 µm, straight to curved, 0–10-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base truncate to short obconically truncate, 1.5–2 µm wide, hila unthickened, not darkened.

Holotype: India: West Bengal: Calcutta, Presidency College, on Celosia argentea, 17 Feb. 1967, A. K. Kar & M. Mandal (K(M) IMI 135869).

Host range and distribution: On Celosia argentea [cristata], Amaranthaceae, Asia (China; India, West Bengal; Japan).

Notes: The conidial width in this species is uniform, but the length rather variable, and relatively short in the type material. The length of conidiophores is also rather variable. Japanese collections of this species were originally wrongly identified and published as Cercospora celosiae (Katsuki 1965), which was clarified in Nakashima et al. (2002).

Pseudocercospora chamissoana R.F. Castaneda & U. Braun, Cryptog. Bot. 1: 51 (1989). (Fig. 85)

Fig. 85
figure85

Pseudocercospora chamissoana (HAL 1651 F, isotype). A. Conidiophores fascicles. B. Conidia. Bar = 10 µm.

Illustration: Castañeda Ruiz & Braun (1989: 47, pl. 3, fig. 18).

Description: Leaf spots amphigenous, subcircular to irregular, 1–10 mm diam, brownish, later pale, margin indefinite or narrow, light to dark brown, formed as mariginal line. Caespituli hypophyllous, punctiform, dark brown to blackish, scattered. Mycelium internal. Stromata substomatal, 20–45 µm diam, brown, composed of swollen hyphal cells, subglobose to angular in outline, 2–5 µm diam. Conidiophores in small to moderately large fascicles, divergent to dense, arising from stromata, through stomata, erect, straight, subcylindrical-conical to geniculate-sinuous, unbranched, 5–25 × 2–5 µm, 0–1-septate, pale brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 5–20 µm long, conidiogenous loci inconspicuous to subdenticulate, but always unthickened and not darkened. Conidia solitary, obclavate-cylindrical to somewhat fusoid, straight to slightly curved, 40–65 × 2–3 µm, indistinctly 3–6-septate, subhyaline to pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base obconically truncate, 1–2 µm wide, hila neither thickened nor darkened.

Holotype: Cuba: Granma: Guisa, Los Corrales, on Chamissoa altissima, Amaranthaceae, 15 Jun. 1987, R. F. Castañeda Ruiz (INIFAT, C87/167). Isotype: HAL 1651 F.

Host range and distribution: Only known from the type collection.

Pseudocercospora cyathulae (Syd.) U. Braun, comb. nov.

MycoBank MB814580 (Fig. 86)

Fig. 86
figure86

Pseudocercospora cyathulae (CUP 39581, holotype). A. Superficial hyphae. B. Solitary conidiophores arising from superficial hyphae. C. Conidiophores. D. Conidia. Bar = 10 µm.

Basionym: Cercospora cyathulae Syd., Ann. Mycol. 35: 239 (1937).

Literature: Chupp (1954: 33), Vasudeva (1963: 95), Kamal (2010: 115–116).

Description: Leaf spots lacking or almost so, on the upper leaf surface only formed as yellowish discolorations, 5–10 mm diam or larger, diffuse, below visible as subcircular brownish patches caused by colonies of the fungus, margin indefinite.

Caespituli hypophyllous, effuse, brownish or dark olivaceous-brown. Mycelium internal and external; superficial hyphae plagiotropous or climbing leaf hairs, forming a dense net or cover, branched, sometimes anastomosing, septate, sterile hyphae narrow, 1–3.5 µm, subhyaline to pale olivaceous or olivaceous-brown, mostly without constrictions at septa, fertile hyphae with conidiophores or conidiogenous cells broader and darker, 3–8 µm wide, swollen cells occasionally to 12 µm diam, olivaceous to medium olivaceous-brown, often with constrictions at septa, thin-walled, smooth. Stromata lacking. Conidiophores solitary, arising from superficial hyphae, lateral or terminal, length indefinite (forming decumbent fertile threats, but differentiation between individual conidiophores and hyphal portions difficult or even impossible), shorter lateral conidiophores arising from decumbent threats about 5–30 µm long, width about 3–8 µm, erect to decumbent, straight, subcylindrical to geniculate-sinuous, unbranched or branched, aseptate to pluriseptate, olivaceous, olivaceous-brown to light brown, thin-walled, smooth; conidiogenous cells integrated, terminal, 10–25 µm long, conidiogenous loci inconspicuous to subconspicuous by being slightly refractive, but unthickened, occasionally visible in front view as minute circle (paracercosporoid) or subdenticulate, 1.5–2 µm diam. Conidia solitary, rarely in short chains, cylindrical to vermiform, shorter conidia sometimes ovoid, obovoid, short cylindrical, broadly fusiform, straight to curved, occasionally sigmoid, 15–100 × (4−)5–8 µm, 0–7-septate, occasionally somewhat constricted at the septa, pale brown, thin-walled, smooth, apex broadly rounded, rarely subtruncate or somewhat attenuated, base rounded to short obconically truncate, 1.5–2 µm wide, hila unthickened, not darkened, sometimes somewhat darker by being refractive.

Holotype: India: Uttarakhand: Dehradun, Mussoorie, Rajpur, Cyathula tomentosa, Amaranthaceae, 18 Sep. 1933, R. N. Tandon 205 (CUP 39581).

Host range and distribution: Only known from the type collection.

Notes: The CUP collection is the only material of this species that could be traced and examined. Chupp (1954) confused this species with Ragnhildiana cyathulae (≡ Passalora cyathulae) and used the wrong citation “Cercospora cyathulae (Stev. & Solh.) Sydow” although Sydow (1937) did not intend to introduce a new combination based on Ragnhildiana cyathulae. He undoubtedly published a new species without any reference to R. cyathulae. The two species are neither conspecific nor congeneric.

Pseudocercospora froelichiae U. Braun & F.O. Freire, Cryptog. Mycol. 25: 230 (2004). (Fig. 87)

Fig. 87
figure87

Pseudocercospora froelichiae (HAL 1779 F, holotype). A. Conidiophores fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustrations: Braun & Freire (2004: 231, fig. 7).

Description: Leaf spots lacking or only with inconspicuous discolorations, yellowish ochraceous, brownish or occasionally purplish, 1–5 mm diam. Colonies formed on the upper leaf surface as sooty patches caused by dense fructification. Mycelium internal. Stromata immersed or somewhat erumpent, 10–50 µm diam, olivaceous-brown, composed of swollen hyphal cells, 2–7 µm diam, walls somewhat thickened. Conidiophores in small to moderately large fascicles, loose to dense, arising from stromata, erumpent, erect, straight, subcylindrical-conical, slightly geniculate-sinuous, unbranched, 10–35 × 2–5 µm, 0–1(− 2)-septate, pale olivaceous to olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci inconspicuous. Conidia solitary, obclavate-cylindrical, 10–80 × 2–3.5 µm, 1–7-septate, pale olivaceous, thin-walled, smooth, apex obtuse or subacute, base short obconically truncate, 1–2 µm wide, hila unthickened, not darkened.

Holotype: Brazil: State of Ceará: Paraipaba City, on Froelichia sp., 6 Aug. 2002, F. O. Freire (HAL 1779 F). Paratype: Brazil: Rio Grande do Norte State: Areia Branca City, on Froelichia sp., 25 Aug. 2003, F. O. Freire (HAL 1780 F).

Host range and distribution: On Froelichia sp., Amaranthaceae, South America (Brazil, State of Cearâ, Rio Grande do Norte State).

Note: Pseudocercospora froelichiae on Froelichia sp. (Amaranthaceae, Gomphrenoideae) is morphologically close to P. chamissoana, described from Cuba on Chamissoa altissima (Amaranthaceae, Amaranthoideae), which differs, however, in forming distinct leaf spots, hypophyllous caespituli and substomatal stromata.

Pseudocercospora globosae (J.M. Yen) Deighton, Mycol. Pap. 140: 144 (1976). (Fig. 88)

Fig. 88
figure88

Pseudocercospora globosae (PC, holotype). A. Conidiophores fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Basionym: Cercospora globosae J.M. Yen, Rev. Mycol. 29: 224 (1964).

Literature: Yen & Lim (1980: 177), Braun & Sivapalan (1999: 14), Crous & Braun (2003: 199).

Illustrations: Yen (1964: 227, fig. 7), Yen & Lim (1980: 239, fig. 42).

Description: Leaf spots at first inconspicuous, later subcircular to somewhat irregular, greyish white to dark grey by abundant fungal colonies. Caespituli amphigenous, effuse to dense, velvety, greyish white. Mycelium internal. Stromata subglobose to somewhat irregular, substomatal to immersed, 10–40 µm diam, yellowish brown. Conidiophores in small to moderately large fascicles, loose to moderately dense, arising from stromata, through stomata or erumpent, straight and subcylindrical-conical to distinctly geniculate-sinuous, unbranched to branched, 10–100 × 1.5–5.5 µm, 0–6-septate, subhyaline to pale yellowish or olivaceous-brown, thin-walled, smooth; conidiogenous cells integrated, terminal, occasionally intercalary, 10–25 µm long, conidiogenous loci inconspicuous to distinctly denticle-like, subcylindrical-conical, apex truncate, 1–2 µm wide, always unthickened and not darkened. Conidia solitary, filiform-subcylindrical, subacicular to narrowly obclavate-subcylindrical, (40−)60–130(− 150) × 1.5–3.5 µm, 2–12-septate, subhyaline to very pale olivaceous or olivaceous-brown, thin-walled, smooth, apex subacute to subobtuse, base truncate to short or long obconically truncate, 1–1.5 µm wide, hila unthickened, not darkened.

Holotype: Singapore: Katong, on Gomphrena globosa, 20 Apr. 1964, S. H. Yen 19 (PC).

Host range and distribution: On Gomphrena (globosa, Gomphrena sp.), Amaranthaceae, Asia (Brunei, Singapore).

Pseudocercospora gomphrenae Goh & W.H. Hsieh, Trans. Mycol. Soc. Republ. China 4(2–3): 8 (1989). (Fig. 89)

Fig. 89
figure89

Pseudocercospora gomphrenae (based on Hsieh & Goh 1990: 19, fig. 8). A. Conidiophore fascicle. B. Conidia. Bar = 10 µm.

Synonym: Cercospora gomphrenae Sawada, Rep. Gov. Res. Inst. Formosa 85: 107 (1943), nom. inval. (Art. 39.1) [type: see Pseudocercospora gomphrenae].

Literature: Hsieh & Goh (1990: 18–19), Guo & Hsieh (1995: 10), Guo et al. (1998: 21).

Illustrations: Hsieh & Goh (1990: 19, fig. 8), Guo & Hsieh (1995: 11, fig. 10), Guo et al. (1998: 21, fig. 10).

Description: Leaf spots amphigenous, subcircular to elliptical, 2–10 mm diam, centre greyish white, margin reddish, on the upper surface with yellowish halo, below olivaceous with dark olivaceous to brown border. Caespituli hypophyllous. Mycelium internal. Stromata lacking. Conidiophores in small fascicles, not more than six, arising from internal hyphae, erect, straight, subcylindrical to somewhat attenuated towards the tip, slightly geniculate-sinuous, unbranched, 20–30 × 3.5–4 µm, 0–2-septate, olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, conidiogenous loci inconspicuous, unthickened, not darkened. Conidia solitary, cylindrical, obclavate-cylindrical to subacicular, straight to curved, 30–90 × 2.5–4 µm, 4–7-septate, subhyaline to very pale olivaceous, thin-walled, smooth, apex obtuse to subacute, base subtruncate to obconically truncate, about 1.5–2 µm wide, hila unthickened, not darkened.

Lectotype (designated here, MycoBank, MBT202805): Taiwan: Taipei, on Gomphrena globosa, 15 Nov. 1925, K. Sawada (NTU-PPE, hb. Sawada). Isolectotype: TNS-F-220432

Host range and distribution: On Gomphrena globosa, Amaranthaceae, Asia (Brunei, China, Iran, Singapore, Taiwan).

Notes: Records of this species from Iran are based on Hedjaroude (1976) and Bakshi et al. (2012).

Pseudocercospora gomphrenae-pulchellae U. Braun et al., Fungal Diversity 6: 28 (2001). (Fig. 90)

Fig. 90
figure90

Pseudocercospora gomphrenae-pulchellae (HAL 1635 F, isotype). A. Conidiophores fascicles. B. Conidiophores. C. Conidia. Bar = 10 µm.

Illustration: Braun et al. (2001: 29, fig. 8).

Description: Leaf spots amphigenous, subcircular to irregular, 1–5(− 8) mm diam, centre pale, yellowish to ochraceous or greyish white, margin narrow, dark reddish brown to blackish brown. Caespituli amphigenous, punctiform, loose to dense, blackish, later greyish white by abundant conidial formation. Mycelium internal; hyphae branched, 1.5–6 µm diam, septate, subhyaline to pale brown, smooth. Stromata substomatal to intraepidermal, 10–100 µm diam, olivaceous-brown. Conidiophores in small to large fascicles, moderately dense, arising from stromata, through stomata or erumpent. erect, straight, subcylindrical or attenuated towards the tip, geniculate-sinuous, unbranched, 10–30 × 2.5–6 µm, 0–1-septate, subhyaline to pale olivaceous, thin-walled, smooth; conidiogenous cells integrated, terminal or conidiophores reduced to conidiogenous cells, 10–25 µm long, conidiogenous loci inconspicuous, occasionally subconspicuous, paracercosporoid, i.e. rim very slightly darkened or thickened, in front view visible as minute circle. Conidia solitary, obclavate-cylindrical, 20–90 × (2−)3–5(− 6) µm, (0−)1–8-septate, subhyaline (with a very pale greenish tinge), smooth, apex obtuse, occasionally subacute, base rounded, truncate to obconically truncate, 1.5–3 µm wide, hila unthickened, not darkened.

Holotype: Argentina: Prov. Buenos Aires: Bahia Blanca, on Gomphrena pulchella, Amaranthaceae, 17 Mar. 2000, R. Delhey 1340 (BB). Isotype: HAL 1635 F.

Host range and distribution: Only known from the type collection.

Note: Pseudocercospora gomphrenae differs from P. gomphrenae-pulchellae in lacking stromata, and P. globosa has smaller stromata, much longer, septate conidiophores and narrower conidia, only 1.5–3.5 µm wide.

Doubtful, excluded and insufficiently known species

Pseudocercospora gomphrenicola Chidd., Sci. & Cult. 22: 511 (1957).

Synonym: Alternaria gomphrenae Togashi, Bull. Imp. Coll. Agric. For. Morioka 9: 6 (1926).

Literature: Kamal (2010: 179).

Type: India: Maharashtra: Pune, on Gomphrena globosa, P. P. Chiddarwar (otherwise not specified).

Note: Type material of this species was not indicated, but based on the original description and illustration of a species with cicatrized conidiogenous cells and rostrate conidia, Kamal (2010) reduced P. gomphrenicola to synonymy with Alternaria gomphrenae.

References

  1. Ahmad S, Iqbal SH, Khalid AN (1997) Fungi of Pakistan. Lahore: Sultan Ahmad Mycological Society of Pakistan.

  2. Alfieri JrSA, Langdon KR, Wehburg C, Kimbrough JW (1984) Index of Plant Diseases on Florida (Revised). Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Bulletin 11: 1–389.

  3. Atkinson GF (1892) Some Cercosporae from Alabama. Journal of the Elisha Mitchel Science Society 8: 33–76.

  4. Bagyanarayana G, Jagadeeswar P, Braun U (1991) Miscellaneous notes on Indian Cercosporae. Mycotaxon 42: 319–326.

  5. Bakalova GG, Borisova TI (2010) Cercosporoid Hyphomycetous Fungi. [Fungi of Bulgaria vol. 7.] Sofia: Institute of Botany, Bulgarian Academy of Sciences.

  6. Bakshi M, Arzanlou M, Babai-Ahari A (2012) Comprehensive check list of cercosporoid fungi from Iran. Plant Pathology & Quarantine 2: 44–55.

  7. Bhartiya HD, Narayan S, Singh AN, Kumari N (2003) Additions to Cercospora from North-Eastern Uttar Pradesh. Indian Phytopathology 56: 270–275.

  8. Bhartiya HD, Singh AN, Kumari N, Singh PN (1998) [“1997”] Foliicolous fungi in the forest flora of North-Eastern Uttar Pradesh. Kavaka 25: 43–51.

  9. Boedijn KB (1961) The genus Cercospora in Indonesia. Nova Hedwigia 3: 411–436.

  10. Braun U (1992) Taxonomic notes on some species of the Cercospora complex. Nova Hedwigia 55: 211–221.

  11. Braun U (1993) Taxonomic notes on some species of the Cercospora complex (III). Mycotaxon 48: 275–298.

  12. Braun U (1995) A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic hyphomycetes). Vol. 1. Eching: IHW-Verlag.

  13. Braun U (1998) A monograph of Cercosporella, Ramularia and allied genera (phytopathogenic hyphomycetes). Vol. 2. Eching: IHW-Verlag.

  14. Braun, U. (1999) Taxonomic notes on some species of the Cercospora complex (V). Schlechtendalia 2: 1–28.

  15. Braun U (2000a) Miscellaneous notes on some micromycetes. Schlechtendalia 5: 31–56.

  16. Braun U (2000b) Annotated list of Cercospora spp. described by C. Spegazzini. Schlechtendalia 5: 57–79.

  17. Braun U (2001) Taxonomic notes on some species of the Cercospora complex (VII). Fungal Diversity 8: 41–71.

  18. Braun U, Castañeda Ruiz RF (1991) Cercospora and allied genera of Cuba (II). Cryptogamic Botany 2–3: 289–297.

  19. Braun U, Crous PW (2005) Additions and corrections to names published in Cercospora and Passalora. Mycotaxon 92: 395–416.

  20. Braun U, Freire FCO (2004) Some cercosporoid hyphomycetes from Brazil — III. Cryptogamie Mycologie 25: 221–244.

  21. Braun U, Freire FCO (2006) Some cercosporoid hyphomycetes from Brazil — IV. Cryptogamie Mycologie 27: 231–248.

  22. Braun U, Hill CF (2002) Some new micromycetes from New Zealand. Mycological Progress 1: 19–30.

  23. Braun U, Mel’nik VA (1997) Cercosporoid fungi from Russia and adjacent countries. Trudy Botanicheskogo Instituta Imeni V.L. Komarova, Rossijskaya Akademiya Nauk St. Petersburg 20: 1–130.

  24. Braun U, Sivapalan A (1999) Cercosporoid hyphomycetes from Brunei. Fungal Diversity 3: 1–27.

  25. Braun U, Urtiaga R (2008) New species and new records of cercosporoid hyphomycetes from Venezuela. Feddes Repertorium 119: 484–506.

  26. Braun U, Urtiaga R (2013) New species and new records of cercosporoid hyphomycetes from Cuba and Venezuela (Part 2). Mycosphere 4: 165–205

  27. Braun U, Bagyanarayana G, Jagadeeswar P (1992) Notes on Indian Cercosporae and allied genera (II). International Journal of Mycology and Lichenology 4: 361–374.

  28. Braun U, Crous PW, Nakashima C (2014) Cercosporoid fungi (Mycosphaerellaceae) 2. Species on monocots (Acoraceae to Xyridaceae, excluding Poaceae). IMA Fungus 5: 203–390.

  29. Braun U, Crous PW, Nakashima C (2015) Cercosporoid fungi (Mycosphaerellaceae) 3. Species on monocots (Poaceae, true grasses). IMA Fungus 6: 25–97.

  30. Braun U, Crous PW, Pons N (2002) Annotated list of Cercospora species (epithets a-b) described by C. Chupp. Feddes Repertorium 113: 112–127.

  31. Braun U, Delhey R, Kiehr M (2001) Notes on some cercosporoid hyphomycetes from Argentina. Fungal Diversity 6: 18–33.

  32. Braun U, Nakashima C, Crous PW (2013) Cercosporoid fungi (Mycosphaerellaceae) 1. Species on other fungi, Pteridophyta and Gymnospermae. IMA Fungus 4: 265–345.

  33. Bridson GDR (2004a) BPH-2, Periodicals with botanical content, constituting a second edition of Botanico-Periodicum-Huntianum. Vol. 1, A-M. Pittsburgh: Hunt Institute for Botanical Documentation.

  34. Bridson GDR (2004b) BPH-2, Periodicals with botanical content, constituting a second edition of Botanico-Periodicum-Huntianum. Vol. 2, N–Z. Pittsburgh: Hunt Institute for Botanical Documentation.

  35. Brummitt RK, Powell CE (1992) Authors of Plant Names. Kew: Royal Botanic Gardens.

  36. Castañeda Ruiz RF, Braun U (1989) Cercospora and allied genera of Cuba (I). Cryptogamic Botany 1: 42–55.

  37. Chaudhary RK, Narayan S, Uphadhyay UC, Rao GP (1995) New species of Cercospora causing foliar diseases in weed flora of sugarcane crop in North-Eastern, U.P. Journal of Living World 2(2): 38–45.

  38. Chi PK (1994) Fungal Diseases of Cultivated Medicinal Plants in Guangdong Province. Guangzhou: Guangdong Academic Press.

  39. Chiddarwar PP (1959) [“1960”] Contributions to our knowledge of the Cercosporae of Bombay State — I. Sydowia 13: 152–163.

  40. Chiddarwar PP (1962) Contributions to our knowledge of the Cercosporae of Bombay State — III. Mycopathologia et Mycologia Applicata 17: 71–81.

  41. Chowdhury S (1956) [“1955”] Notes on fungi of Assam, I. Lloydia 18: 82–87.

  42. Chupp C (1954) A Monograph of the fungus genus Cercospora. Ithaca, NY: C. Chupp.

  43. Clement W Arikake M, Sweeney P, Edwards EJ, Donoghue MJ (2014) A chloroplast tree for Viburnum (Adoxaceae) and its implication for phylogenetic classification and character evolution. American Journal of Botany 101: 1029–1049.

  44. Crous PW, Braun U (1996). Cercosporoid fungi from South Africa. Mycotaxon 57: 233–321.

  45. Crous PW, Braun U (2001) A reassessment of the Cercospora species described by C. Chupp: specimens deposited at BPI, Maryland, U.S.A. Mycotaxon 78: 327–343.

  46. Crous PW, Braun U (2003) Mycosphaerella and its Anamorphs: 1. Names published in Cercospora and Passalora. [CBS Biodiversity Series no. 1]. Utrecht: Centraalbureau voor Schimmelcultures.

  47. Crous PW, Braun U, Hunter GC, Wngfield MJ, Verkley GJM, et al. (2013) Phylogenetic lineages in Pseudocercospora. Studies in Mycology 75: 37–114.

  48. Deighton FC (1974) Studies on Cercospora and allied genera. V Mycovellosiella Rangel, and a new species of Ramulariopsis. Mycological Papers 137: 1–75.

  49. Deighton FC (1976) Studies on Cercospora and allied genera. VI. Pseudocercospora Speg., Pantospora Cif. and Cercoseptoria Petr. Mycological Papers 140: 1–168.

  50. Deighton FC (1979) Studies on Cercospora and allied genera. VII. New species and redispositions. Mycological Papers 144: 1–56.

  51. Deighton FC (1987) New species of Pseudocercospora and Mycovellosiella, and new combinations into Pseudocercospora and Phaeoramularia. Transactions of the British Mycological Society 88: 365–391.

  52. Dubey RK, Rai AN, Verma NK, Thakur RS (2011) Three novel additions to fungal genus Cercospora Fres. from Central India. Journal of Mycology and Plant Pathology 41: 512–517.

  53. Egorova LN (2007) Additions to the biota of anamorphic fungi of the Zeysky Reserve (Amur region, Russia). Mikologiya i Fitopatologiya 41: 202–207.

  54. Ellis MB (1971) Dematiaceous Hyphomycetes. Kew: Commonwealth Mycological Institute.

  55. Ellis MB (1976) More Dematiaceous Hyphomycetes. Kew: Commonwealth Mycological Institute.

  56. Ferraris T (1910) Flora italica cryptogama. Pars I. Fungi, Hyphales. Rocca S. Casciano: Stabilimeto tipografico Cappalli.

  57. Fuckel KWGL (1863) Fungi rhenani exsiccati a L. Fuckel collecti. Fasc. I–IV. 1863. Hedwigia 2: 132–136.

  58. Fuckel KWGL (1870) [“1869”] Symbolae mycologicae: Beitrag zur Kenntnis der rheinischen Pilze. Jahrbücher des Nassauischen Vereins für Naturkunde 23–24: 1–459.

  59. Gola G (1930) L’Erbario Micologoco di P. A. Saccardo. Catalogo. Padova: Tipografia Editrice Antoniana.

  60. González Fragoso DR (1927) Estudio sistemático de los hifales de la flora Española conocidos hasta esta fecha. Memorias de la Real Academia de Ciencias Exactas, Físicas y Naturales de Madrid, 2 Serie, 6: 1–377.

  61. Govindu HC, Thirumalachar MJ (1954) Notes on some Indian Cercosporae. IV. Sydowia 8: 221–230.

  62. Govindu HC, Thirumalachar MJ (1955) Notes on some Indian Cercosporae. VI. Sydowia 9: 221–228.

  63. Govindu HC, Thirumalachar MJ (1957) Notes on some Indian Cercosporae. VIII. Sydowia 10: 271–277.

  64. Groenewald JZ, Nakashima C, Nishikawa J, Shin, HD, Park JH, et al. (2013) Species concepts in Cercospora: spotting the weeds among the roses. Studies in Mycology 75: 115–170.

  65. Guo YL (1995) [“1994”] Four new species of Pseudocercospora. Mycosystema 7: 119–127.

  66. Guo YL (1996) [“1995–1996”] Cercospora and allied genera and species recorded from the Qinling Mountains. Mycosystema 8–9: 89–102.

  67. Guo YL (2001) New species and new records of fungi from tropical China: Hyphomycetes. Mycotaxon 77: 343–348.

  68. Guo YL (2012) Studies on Cercospora and allied genera in China XV. Mycosystema 31: 159–164.

  69. Guo YL, Hsieh WH (1995) The genus Pseudocercospora in China. Mycosystema Monographicum Series 2: 1–388.

  70. Guo YL, Jiang Y (2000) Studies on Cercospora and allied genera in China I. Mycotaxon 74: 257–266.

  71. Guo YL, Liu XJ (1992) Studies on the genus Pseudocercospora in China VI. Mycosystema 5: 99–108.

  72. Guo YL, Liu XJ, Hsieh WH (1998) Pseudocercospora. [Flora Fungorum Sinicorum vol. 9.] Beijing: Science Press.

  73. Guo YL, Liu XJ, Hsieh WH (2003) Mycovellosiella, Passalora, Phaeoramularia. [Flora Fungorum Sinicorum vol. 20.] Beijing: Science Press.

  74. Guo YL, Liu XJ, Hsieh WH (2005) Cercospora. [Flora Fungorum Sinicorum vol. 24.] Beijing: Science Press.

  75. Gupta BK, Kamal (1987) Some new species of Pseudocercospora from India. In Perspectives in Mycological Research — I. Prof. G.P. Agarwal Festschrift Volume (Hasija SK, Rayak RC, Singh SM eds): 19–33. New Delhi: Today & Tomorrow’s Printers and Publishers.

  76. Gupta C, Abbasi P, Kamal (1987) Some new foliicolous hyphomycetes from North-Eastern U.P In Perspectives in Mycological Research — I. Prof. G.PAgarwal Festschrift Volume (Hasija SK, Rayak RC, Singh SM eds): 7–17. New Delhi: Today & Tomorrow’s Printers and Publishers.

  77. Hesami S, Khodaparast SA, Zare R (2011) New reports on Cercospora and Cercospora-like fungi from Guilan Province, Iran. Iranian Journal of Plant Pathology 47: 131–132.

  78. Hsieh WH, Goh TK (1990) Cercospora and similar Fungi from Taiwan. Taipei: Maw Chang Book Company.

  79. Ibrahim FM, El Nur Elamin (1974) A quantitative morphological classification of thirty species of Cercospora. Mycopathologia et Mycologia Applicata 52: 141–146.

  80. Jaganathan T, Palaniswami A, Narayanaswami P (1972) A new leaf spot disease of Crossandra. The Madras Agricultural Journal 59: 671–672.

  81. Jage H, Braun U (2004) Neufunde pflanzenbewohnender Mikromyceten aus der Bundesrepublik Deutschland. Feddes Repertorium 115: 56–61.

  82. Kamal (2010) Cercosporoid Fungi of India. Dehra Dun: Bishen Singh Mahendra Pal Singh.

  83. Kar AK, Mandal M (1969) New Cercospora spp. from West Bengal. Transactions of the British Mycological Society 53: 337–360.

  84. Kar AK, Mandal M (1970) New Cercospora spp. from West Bengal. II. Transactions of the British Mycological Society 54: 423–433.

  85. Karan D, Manoharachary C (1978) [“1976”] Notes on microfungi from Andhra Pradesh — VII Some phytopathogenic fungi from Hyderabad. The Botanique (Nagpur) 7: 157–162.

  86. Katsuki S (1965) Cercosporae of Japan. Transactions of the Mycological Society of Japan, Extra Issue 1: 1–100.

  87. Keissler K, Lohwag H (1937) Fungi. In Handel-Mazzetti H. (ed.): Symbolae sinicae 2: 1–79. Vienna: J. Springer.

  88. Kim JD, Shin HD (1999) Taxonomic studies on Cercospora and allied genera in Korea (X). Korean Journal of Mycology 27: 220–230.

  89. Kirk PM (1999) Stigmina tinea. CMI Descriptions of Pathogenic Fungi and Bacteria 1400: 1–2.

  90. Kobayashi T (2007) Index of fungi inhabiting woody plants in Japan — Host, Distribution and Literature. Japan. Zenkoku-Noson-Kyoiku Kyokai, Publishing.

  91. Kobayashi T, Nakashima C, Nishijima T (2002) Addition and re-examination of Japanese species belonging to the genus Cercospora and allied genera. V Collections from the Nansei Islands (2). Mycoscience 43: 219–227.

  92. Lindau G (1910) Fungi imperfecti: Hyphomycetes (zweite Hälfte), Dematiaceae (Phaeophragmiae bis Phaeostaurosporae, Stilbaceae, Tuberculariaceae, sowie Nachträge, Nährpflanzenverzeichnis und Register. [Rabenhorst’s Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz 1 (9).] Leipzig: Verlag von E. Kummer.

  93. Liu XJ, Guo YL (1982) Studies on some species of the genus Phaeoramularia in China. Acta Phytopathologica Sinica 12(4): 1–15.

  94. Liu XJ, Guo YL (1987) [“1986”] Three hyphomycete species parasitic on Actinidia plants in China. Acta Mycologica Sinica, Supplement 1: 353–358.

  95. Meeboon J, Hidayat I, To-anun C (2007a) Cercosporoid fungi from Thailand 3. Two new species of Passalora and six new records of Cercospora species. Mycotaxon 102: 139–145.

  96. Meeboon J, Hidayat I, To-anun C (2007b) An annotated list of cercosporoid fungi in Northern Thailand. Journal of Agricultural Technology 3: 51–63.

  97. Mendes MAS, da Silva VL, Dianese JC, Ferreira MASV, dos Santos CEN, et al. (1998) Fungos em Plantas no Brasil. Brasília, DF: Servição de Informação — SPI.

  98. Minter DW, Rodriguez Hernández M, Mena Portales J (2001) Fungi of the Carribean: an annotated checklist. Isleworth: PDMS Publishing.

  99. Muntañola M (1960) Algunos Hyphomycetes criticos. Notas y descripciones. Lilloa 30: 165–232.

  100. Nakashima C, Oetari A, Kanti A, Saraswati R, Wdyastuti Y, Ando K (2010) New species and newly recorded species of Cercospora and allied genera from Indonesia. Mycosphere 1: 315–323.

  101. Nakashima C, Tanda S, Kobayashi T (2002) Addition and re-examination of Japanese species belonging to the genus Cercospora and allied genera. IV Newly recorded species from Japan (1). Mycoscience 43: 95–102.

  102. Nguanhom J, Cheewangkoon R, Groenewald JZ, Braun U, To-Anun C, Crous PW (2015) Taxonomy and phylogeny of Cercospora spp. from Northern Thailand. Phytotaxa 233(1): 027–048.

  103. Patil MS (1975) Some Cercospora species from Kolhapur — II. Botanique 6: 219–226.

  104. Patil MS (1978) Some Cercospora species from Kolhapur — IV. Botanique 8: 69–74.

  105. Patwardhan PG, Pande AK (1970) [“1969”] Some Fungi-Imperfecti from Maharashtra. Sydowia 23: 95–101.

  106. Pavgi MS, Singh RA (1964) Parasitic fungi from North India. Mycopathologia et Mycologia Applicata 23: 188–196.

  107. Pavgi MS, Singh RA (1965) Parasitic fungi from North India — V. Mycopathologia et Mycologia Applicata 27: 89–96.

  108. Phengsintham P, Braun U, McKenzie EHC, Chukeatirote E, Cai L, Hyde KD (2013) Monograph of cercosporoid fungi from Thailand. Plant Pathology & Quarantine 3: 19–90.

  109. Pirnia M, Zare R, Zamanizadeh HR, Khodaparast A (2010) Contribution to the identification of Cercospora species in Iran. Rostaniha 11: 183–189.

  110. Ponnappa KM (1968) Some interesting fungi-II. Cercospora hygrophilae sp. nov. and Stenella plectroniae sp. nov. Proceedings of the Indian Academy of Sciences, sect. B, 67: 31–34.

  111. Pons N, Sutton BC (1988) Cercospora and similar fungi on yams (Dioscorea spp.). Mycological Papers 160: 1–78.

  112. Raghu Ram M, Mallaiah KV (1996) New species of Mycovellosiella and Cercospora from India. Mycological Research 100: 295–296.

  113. Rai AN, Kamal (1987) New Cercospora species from India. Transactions of the British Mycological Society 89: 124–126.

  114. Rao GP, Filho AB, Magarey RC, Autrey LJC (1999) Sugarcane Pathology. Volume I: Fungal Diseases. Enfield, NH: Science Publisher Inc.

  115. Ray WW (1941) Notes on Oklahoma Cercosporae. Mycologia 33: 174–177.

  116. Saccardo PA (1886) Sylloge Fungorum omnium hucusque cognitum. Vol. 4. Padova: P.A. Saccardo.

  117. Saccardo PA (1892) Sylloge Fungorum omnium hucusque cognitum. Vol. 10. Padova: P.A. Saccardo.

  118. Saccardo PA (1895) Sylloge Fungorum omnium hucusque cognitum. Vol. 11. Padova: P.A. Saccardo.

  119. Saccardo PA (1906) Sylloge Fungorum omnium hucusque cognitum, Vol. 18. Supplementum Universal Pars VII. [Saccardo PA, Saccardo D, eds]. Padova: P.A. Saccardo.

  120. Saccardo PA (1913) Sylloge Fungorum omnium hucusque cognitum, Vol. 22. [Saccardo PA, Trotter A, eds]. Padova: P.A. Saccardo.

  121. Saccardo PA (1972) Sylloge Fungorum omnium hucusque cognitum. Vol. 26 [Trotter A, Cash K, eds]. New York: Johnson.

  122. Shin HD, Kim JD (2001) Cercospora and allied genera from Korea. Plant Pathogens of Korea 7: 1–303.

  123. Shivas RG, Suyoko S, Raga N, Hyde KD (1996) Some disease-associated microorganisms on plants in Irian Jaya, Indonesia. Australasian Plant Pathology 25: 36–49.

  124. Simmons EG (1995) Alternaria themes and variations (112–144). Mycotaxon 55: 55–163.

  125. Singh SM (1976) Some Deuteromycetes from Balaghat, Madhya Pradesh. Indian Phytopathology 29: 17–19.

  126. Singh PN, Singh SK, Tripathi SC (1996) New species of Pseudocercospora causing leaf spots of forest plants of Nepal. Mycological Research 100: 1129–1132.

  127. Solheim WG, Stevens F (1931) Cercospora studies: II. Some tropical Cercosporae. Mycologia 23: 365–405.

  128. Srivastava RK, Narayan S, Kumari N (2001) New Cercospora spp. from north-eastern Uttar Pradesh. Indian Phytopathology 54: 102–107.

  129. Srivastava RK, Srivastava AK, Kamal (1995) [“1992–1993”] An appraisal of Indian species of Cercospora Fres. sensu stricto: some new species and new records. Kavaka 20/21: 37–47.

  130. Stevenson JA (1975) Fungi of Puerto Rico and the American Virgin Islands. Contributions from the Reed Herbarium 23: 1–742.

  131. Sutton BC, Pons N (1980) Notes on the original species of Cercosporina. Mycotaxon 12: 201–218.

  132. Swiderska-Burek U, Mutenko W (2014) New records of cercosporoid fungi from Poland. Mycotaxon 128: 55–62.

  133. Sydow H, Mitter JH, Tandon RN (1937) Fungi indici — III. Annales Mycologici 35: 222–243.

  134. Tai FL (1948) Cercosporae of China II. Lloydia 11: 36–56.

  135. Tai FL (1979) Sylloge Fungorum Sinicorum. Beijing: Academia Sinica, Science Press.

  136. Thaung MM (1984) Some fungi of Cercospora from Burma. Mycotaxon 19: 425–452.

  137. Thirumalachar MJ, Chupp C (1948): Notes on some Cercosporae from India. Mycologia 40: 352–362.

  138. Thirumalachar MJ, Govindu HC (1953) Notes on some Cercosporae of India — II. Sydowia 7: 45–49.

  139. To-anun C, Hidayat I, Meeboon J (2011) Genus Cercospora in Thailand: taxonomy and phylogeny (with a dichotomous key to species). Plant Pathology & Quarantine 1: 11–87.

  140. Vassiljevsky NI, Karakulin BP (1937) Parazitnye nesovershennye griby. Vol. 1. Gifomicety. Moscow, Leningrad: Izdatekl’stvo Akademii Nauk SSSR.

  141. Vasudeva RS (1963) Indian Cercosporae. New Delhi: Indian Council of Agricultural Research.

  142. Verma RK, Kamal (1991) Some new species of Pseudocercospora. Indian Phytopathology 44: 440–447.

  143. Winkworth RC, Donoghue MJ (2005) Viburnum phylogeny based on combined molecular data: implications for taxonomy and biogeography. American Journal of Botany 92: 653–666.

  144. Yen JM (1964) Étude sur les champignons parasites du Sud-Est Asiatique. I: Première note sur quelques nouvelles espèces de Cercospora de Singapour. Revue de Mycologique 29: 209–240.

  145. Yen JM (1965) Étude sur les champignons parasites du Sud-Est Asiatique. III: Deuxième notes su les nouvelles espèces de Cercospora de Singapour. Revue de Mycologie 30: 166–204

  146. Yen JM (1967) Étude sur les champignons parasites du Sud-Est Asiatique. VII. Quatrieme note sur quelques Cercospora et Stenella de Singapour (Malaisie). Revue de Mycologie 32: 177–202.

  147. Yen JM (1977) Étude sur les champignons parasites du Sud-Est Asiatique. XXVI. Les Cercospora de Formose II. Bulletin Trimestriel de la Société Mycologique de France 93: 145–164.

  148. Yen JM (1978) Étude sur les champignons parasites du Sud-Est Asiatique. XXX. Les Cercospora de Formose III. Bulletin Trimestriel de la Société Mycologique de France 94: 49–59.

  149. Yen JM (1983) Studies on parasitic fungi from South East Asia, 49. Parasitic fungi from Malaysia, 25. Semipseudocercospora gen. nov. Mycotaxon 17: 361–363.

  150. Yen JM, Lim G (1973) Étude sur les champignons de Sud-Est asiatique. XX. Dixième note sur les Cercospora de Malaisie. Cahiers du Pacifique 17: 95–144.

  151. Yen JM, Lim G (1980) Cercospora and allied genera of Singapore and the Malay Peninsula. Gardens’ Bulletin, Singapore 33: 151–263.

  152. Yen JM, Kar AK, Das BK (1982) Studies on hyphomycetes from West Bengal, India, I. Cercospora and allied genera of West Bengal, 1. Mycotaxon 16: 35–57.

Download references

Acknowledgements

We are very grateful to the directors and curators of B, BPI, CUP, DAOM, FH, HBG, HMAS, ILL, ILLS, IUM, K, L, LE, LEP, LPS, MA, NIAES, NTU, NY, NYS, PAD, PC, PDD, RREM, S, TFM, TNS and WSP for loaning type material and other collections in their keeping during the course of monographic studies of cercosporoids fungi on the hosts of the families treated in part 4.

Author information

Correspondence to Uwe Braun.

Rights and permissions

This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Braun, U., Crous, P.W. & Nakashima, C. Cercosporoid fungi (Mycosphaerellaceae) 4. Species on dicots (Acanthaceae to Amaranthaceae). IMA Fungus 6, 373–469 (2015) doi:10.5598/imafungus.2015.06.02.09

Download citation

Key words

  • Ascomycota
  • Cercospora s. lat.
  • hyphomycetes
  • taxonomy