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The genus Tremella (Basidiomycota, Tremellales) in Finland

January 2008

January 2008
45(6):401-434

Authors:

Finnish Environment Institute

Sixteen species of Tremella Pers. are currently known from Finland. Fifteen of them are illustrated and described, and a key to all Finnish species is given. Tremella cetrariicola Diederich & Coppins, T. cladoniae Diederich & M.S. Christ., T. giraffa Chee J. Chen, T. globispora D.A. Reid, T.phaeophysciae Diederich & M.S. Christ., T. polyporina D.A. Reid and T. ramalinae Diederich are reported as new to Finland.

Tremella cetrariicola (a–d from Hyvärinen 2005, e–f from Hirvenoja 1951). — a and e: Spores, of which one forms a secondary spore and one germinates by a germ tube. — b and f: Basidia at different stages of development. — c: Hyphae with haustoria. — d: conidia-like cells.

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. Spore dimensions (µm) of selected specimens except for Tremella cladoniae. Specimens are marked with collector's name and collecting number or year; herbarium label is included if needed to separate specimens of the same species. n = number of measured spores. The values set in boldface include at least 90% of the spores.

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Distribution of Tremella cetrariicola () and T. encephala () in Finland. The lines show the boundaries of the biogeographical provinces.

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. Substrates of Tremella encephala, T. foliacea and T. mesenterica.

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+21

Tremella encephala (a–e from Pippola 456, f from Jäppinen 1987). — a and f: Spores, of which some germinate by a germ tube or form secondary spores. — b: Basidia. — c: Hyphae with haustoria, of which one is attached to the host hyphae (arrow). — d: Swollen cells. — e: Hyphidia.

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Ann. Bot. Fennici 45: 401–434 ISSN 0003-3847 (print) ISSN 1797-2442 (online)

Helsinki 30 December 2008 © Finnish Zoological and Botanical Publishing Board 2008

The genus Tremella (Basidiomycota, Tremellales) in

Finland

Emilia Pippola1,* & Heikki Kotiranta2

1) Department of Biology, P.O. Box 3000, FI-90014 University of Oulu, Finland (*corresponding

author’s e-mail: epippola@paju.oulu.ﬁ)

2) Finnish Environment Institute, Research Department, P.O. Box 140, FI-00251 Helsinki, Finland

(heikki.kotiranta@ymparisto.ﬁ)

Received 19 June 2007, revised version received 26 Sep. 2007, accepted 27 Sep. 2007

Pippola, E. & Kotiranta, H. 2008: The genus Tremella (Basidiomycota, Tremellales) in Finland.

— Ann. Bot. Fennici 45: 401–434.

Sixteen species of Tremella Pers. are currently known from Finland. Fifteen of them

are illustrated and described, and a key to all Finnish species is given. Tremella

cetrariicola Diederich & Coppins, T. cladoniae Diederich & M.S. Christ., T. giraffa

Chee J. Chen, T. globispora D.A. Reid, T. phaeophysciae Diederich & M.S. Christ., T.

polyporina D.A. Reid and T. ramalinae Diederich are reported as new to Finland.

Key words: Basidiomycota, fungicolous, heterobasidioid, lichenicolous, parasitism,

taxonomy, Tremella, Tremellales

Introduction

Heterobasidioid fungi are poorly known in Fin-

land. They have largely been overlooked since

the early studies of P. A. Karsten (1834–1917),

except for fragmentary notes in the literature

(e.g., Laurila 1939, Lowy 1960, Ryman & Hol-

måsen 1987, Ohenoja 1996, Hansen & Knudsen

1997, Salo et al. 2006) and studies of resupinate

heterobasidiomycetes (Kotiranta & Saarenoksa

1993, 2005). As a result there is not even much

data on basic species composition. When we

began the study of the genus Tremella, only

nine species had been reported in Finland (Tor-

kelsen 1997, Kotiranta & Saarenoksa 2000). The

number of species was low as compared with

the 25 Tremella species reported in the Nordic

countries, and the distribution and abundance of

the species were insufciently known (Torkelsen

1997). The species were also commonly confused

with other heterobasidioid fungi, especially those

in the genus Exidia and in the order Dacrymyc-

etales. The former group is easily distinguished

from Tremella species by their allantoid spores

and the latter group by their forked basidia.

Material and methods

Approximately 600 Tremella specimens were

examined for the present study. The material is

preserved in the herbaria H, JYV, K, KUO, O,

OULU, S, TUR, UPS and/or in the personal her-

barium of Heikki Kotiranta (H.K.).

Data on the distribution, abundance and ecol-

ogy of Tremella species in Finland are based on

these collections. For each species, except T. cla-

doniae which was added at the last moment, seven

to nine specimens were selected for accurate

measurements. Where less than seven specimens

402 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

exist, all were included. At least thirty spores per

specimen were measured, where present. None

of the measurements derives from spore prints.

Basidia measurements do not include stalks. All

measurements were made using ¥1000 or ¥1500

magnication and oil immersion. An eyepiece

scale bar with 1 µm grid was used, and dimen-

sions were estimated subjectively with an accu-

racy of 0.1 µm. The basic mountant medium used

was 5% KOH. The spore measurements for each

specimen are given in Table 1.

In the table, text and illustrations selected

specimens are marked with the collector’s name

and collecting number or year; the herbarium label

data is included if needed to separate specimens

of the same species. The following abbreviations

are used in the table and text: L = mean length, W

= mean width, Q = L/W , Q* = mean L/W ratio,

Table 1. Spore dimensions (µm) of selected specimens except for Tremella cladoniae. Specimens are marked with

collector’s name and collecting number or year; herbarium label is included if needed to separate specimens of the

same species. n = number of measured spores. The values set in boldface include at least 90% of the spores.

Species/specimen n Spore length (L) Spore width (W ) L/W

Range Mean Range Mean Range Mean

(Q) (Q*)

T. cetrariicola

Hirvenoja 1951 30 (5.2–)5.8–9.6(–9.6) 6.5 (3.8–)4.0–7.8(–8.2) 5.3 1.0–1.5 1.3

Linkola 1909 30 (5.9–)6.1–10.4(–10.4) 7.7 (5.5–)5.7–9.6(–9.8) 6.9 0.9–1.9 1.1

Hyvärinen 2006 30 (4.2–)5.8–8.2(–8.3) 6.3 (3.9–)3.9–6.9(–7.7) 5.2 1.0–1.7 1.2

Hyvärinen 2005 30 (6.1–)6.1–8.4(–9.6) 7.5 (3.9–)4.4–6.5(–6.6) 5.8 1.1–1.7 1.3

T. encephala

Haikonen 23835 31 (7.6–)7.6–11.1(–11.7) 9.4 (6.3–)7.4–9.6(–9.8) 8.5 1.0–1.3 1.1

Karsten 1881 30 (6.5–)7.4–11.6(–11.6) 9.1 (6.0–)6.2–9.9(–10.4) 8.4 1.0–1.3 1.1

Tiensuu 1962 31 (6.4–)7.6–11.6(–12.0) 9.3 (5.9–)7.1–9.8(–10.4) 8.3 0.9–1.3 1.1

Pippola 456 30 (6.0–)6.0–9.8(–9.9) 7.8 (5.0–)5.9–9.8(–9.8) 7.5 0.9–1.2 1.1

Jäppinen 1987 30 (7.6–)7.8–10.1(–10.2) 9.2 (6.0–)6.5–9.2(–9.6) 8.0 1.0–1.3 1.2

Metsänheimo 1984 30 (6.5–)7.8–10.4(–11.8) 9.1 (6.2–)6.4–9.9(–11.6) 8.2 1.0–1.4 1.1

Ohenoja 1977 30 (7.4–)7.9–11.2(–11.2) 9.5 (6.5–)7.4–10.0(–10.1) 8.8 1.0–1.2 1.1

T. foliacea

Stenlid 668 30 6.2–8.2(–8.6) 7.4 (5.8–)5.9–7.9(–8.0) 6.6 1.0–1.3 1.1

Oittinen 1966 30 (8.0–)8.1–10.3(–10.5) 9.3 7.2–9.6(–9.8) 8.4 1.0–1.2 1.1

Malmström 39 30 (6.0–)6.2–9.4(–9.5) 8.0 (5.8–)6.0–8.2(–9.6) 7.1 1.0–1.3 1.1

Kankainen 1966 30 (6.3–)6.9–9.9(–10.0) 8.5 5.9–9.2(–9.8) 7.7 1.0–1.3 1.1

Koskela 1970 30 (6.4–)7.3–10.1(–11.4) 8.3 (5.8–)6.0–9.4(–9.8) 7.6 0.9–1.3 1.1

Ohenoja & Ohenoja 1972 30 (7.4–)7.6–9.7(–9.8) 8.1 (6.0–)6.1–8.2(–8.5) 7.4 1.0–1.5 1.1

Pippola 376 30 (6.0–)6.1–8.3(–8.4) 7.5 (4.5–)5.7–7.6(–7.9) 6.3 1.0–1.4 1.2

T. giraffa

Alanko 132586 31 (5.7–)6.1–8.2(–8.3) 7.3 (5.0–)5.5–7.8(–8.0) 6.6 0.9–1.4 1.1

Alanko 132671 30 (6.2–)7.0–9.1(–9.9) 8.0 (6.0–)6.1–9.7(–9.8) 6.6 0.9–1.3 1.1

T. globispora

Söderholm 2752 30 (5.9–)6.0–9.2(–9.6) 7.5 (5.9–)6.1–9.6(–9.7) 7.9 0.8–1.1 1.0

Karsten 1867 30 (5.8–)7.4–10.2 8.3 (5.4–)6.9–11.6(–12.0) 9.1 0.8–1.1 0.9

Karsten 1862 30 (5.4–)5.8–9.2(–9.8) 7.0 5.9–9.7(–9.9) 7.3 0.8–1.2 1.0

T. hypocenomycis

Leppälä 1947 5 4.6–7.8 6.3 4.0–8.1 6.2 0.9–1.2 1.0

Norrlin 1867 15 5.4–7.3 6.4 5.3–7.6 6.2 0.9–1.2 1.0

T. hypogymniae

Häyrén 1919 30 (4.5–)5.1–7.9(–8.0) 6.5 (4.9–)5.1–8.3(–8.6) 7.1 0.7–1.1 0.9

Takala 73 32 (5.1–)5.3–8.2(–8.2) 6.8 (6.1–)6.2–9.7(–9.8) 8.0 0.6–1.2 0.9

Takala 82 30 5.6–7.8(–8.1) 6.6 (5.8–)5.9–8.6(–9.5) 7.6 0.7–1.3 0.9

Takala 3 30 (4.9–)5.5–7.6(–7.7) 6.4 (5.5–)5.9–8.1(–8.3) 7.4 0.7–1.0 0.9

Räsänen 1936 18 4.1–7.9 6.1 5.5–8.2 7.1 0.7–1.0 0.9

Linkola 1930 30 (4.2–)5.0–7.3(–7.9) 5.9 (5.3–)5.7–7.8(–8.2) 6.8 0.7–1.0 0.9

Takala 103 9 5.9–8.3 6.4 6.3–10.0 8.6 0.7–1.0 0.8

continued

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 403

Table 1. Continued.

Species/specimen n Spore length (L) Spore width (W) L/W

Range Mean Range Mean Range Mean

(Q) (Q*)

T. indecorata

Haikonen 1866 30 (4.9–)5.6–8.0(–9.8) 7.0 (5.7–)5.9–9.2(–9.4) 7.2 0.8–1.2 1.0

Söderholm 2686 30 (5.9–)6.0–9.8(–9.9) 7.8 (5.9–)6.1–10.0(–11.2) 8.0 0.9–1.3 1.0

Karsten 1865 30 (5.5–)5.6–8.8(–9.0) 7.1 (5.9–)5.9–9.8(–9.9) 7.6 0.8–1.0 0.9

Pippola 80 30 (5.9–)6.0–8.3(–9.4) 7.4 (5.7–)5.9–9.8(–10.9) 7.7 0.8–1.1 1.0

Pippola 115 30 (5.9–)6.0–9.4(–9.8) 7.7 (5.9–)6.2–9.8 7.8 0.8–1.2 1.0

T. karstenii

Karsten 30 (2.7–)3.6–5.9(–7.6) 4.5 (3.3–)3.9–6.0(–6.1) 4.8 0.7–1.4 0.9

Fagerström 1976 30 (3.9–)4.0–6.4(–6.5) 5.4 (3.9–)4.3–7.1(–7.5) 5.5 0.8–1.3 1.0

Holm & Holm 1969 30 (3.7–)4.0–5.7(–6.1) 4.6 (4.2–)4.5–6.3(–6.5) 5.6 0.7–1.0 0.8

Karsten 1869 30 (3.2–)3.3–5.2(–5.9) 3.9 3.4–5.2(–5.9) 4.1 0.8–1.1 0.9

Pippola 449 30 3.7–5.9 4.3 (3.7–)3.8–5.8(–5.9) 4.5 0.8–1.3 1.0

Torkelsen 106/78 30 3.9–5.6(–6.1) 4.4 (3.0–)3.8–5.9(–6.5) 4.9 0.7–1.3 0.9

Pippola 231 30 (2.5–)3.9–6.0 4.7 (3.7–)3.9–6.3(–6.7) 5.1 0.7–1.1 0.9

T. mesenterica

Pippola 282 30 (8.0–)8.8–13.1(–13.9) 11.0 7.8–13.3(–13.4) 10.1 0.9–1.3 1.1

Korhonen 6307 30 (9.5–)9.6–13.7(–14.0) 11.7 (7.7–)8.1–11.7(–11.8) 9.8 1.0–1.4 1.2

Haikonen 1942 30 (9.2–)9.8–15.0(–15.6) 12.2 (7.4–)7.8–12.1(–12.9) 9.6 1.0–1.6 1.3

Särkkä 169 30 (9.9–)10.1–13.7(–15.5) 11.9 (7.5–)7.9–11.7(–14.0) 9.8 1.0–1.6 1.2

Ulvinen 1986 30 (11.7–)11.8–15.6(–17.4) 13.8 (10.2–)10.4–14.9(–15.6) 12.4 0.9–1.4 1.1

Pippola 439 30 (9.6–)10.1–14.4(–15.6) 12.9 9.6–13.1(–13.7) 11.7 1.0–1.2 1.1

Kallio 1960 30 9.6–14.5(–15.6) 12.0 (7.6–)7.8–13.5(–13.7) 10.1 1.0–1.4 1.2

T. mycetophiloides

Kotiranta 11860 0 − − − − − −

T. obscura

Pippola 292 1 8.2 8.2 7.8 7.8 1.1 1.1

Laurila 1936 30 (6.1–)6.3–9.8(–10.4) 8.0 (6.0–)6.1–9.6(–9.8) 7.7 0.9–1.2 1.0

Ulvinen 1972 34 (4.3–)4.5–8.0(–8.2) 6.3 3.9–7.6(–7.8) 4.8 1.1–1.7 1.3

Pippola 394 30 (5.9–)6.0–8.2(–8.4) 7.3 (5.4–)5.5–7.5(–7.7) 6.6 1.0–1.3 1.1

T. phaeophysciae

Laine 1989 30 (4.3–)4.9 –7.0(–7.1) 6.0 (5.3–)5.4–8.0(–8.9) 6.8 0.8–1.0 0.9

Hinneri 1970 19 5.7–7.8 6.7 5.9–8.6 7.3 0.8–1.0 0.9

Hausen 1925 30 (4.2–)4.9–6.9(–7.3) 6.0 5.1–8.0(–8.2) 6.8 0.7–1.1 0.9

Hausen 1927

(OULU F073618) 30 4.3–7.6 6.1 (4.5–)4.6–7.8(–8.4) 6.6 0.7–1.1 0.9

Hausen 1927

(OULU F073615) 30 (4.3–)5.1–7.9(–8.0) 6.2 5.9–7.9(–9.4) 6.9 0.7–1.0 0.9

Räsänen 1935 9 5.7–7.8 6.3 6.2–8.0 7.0 0.8–1.0 0.9

Räsänen 1918 2 5.8–6.9 6.4 6.3–7.1 6.7 0.8–1.1 1.0

Räsänen 1920 30 (4.9–)5.0–7.0(–7.2) 6.1 (5.3–)5.8–8.1 7.1 0.7–1.1 0.9

Leppälä 1948 30 (5.2–)5.7–8.1(–9.1) 6.9 (6.2–)6.3–8.4(–9.3) 7.5 0.7–1.1 0.9

T. polyporina

Kiema 176 30 (4.5–)5.1–6.0(–6.3) 5.6 4.4–5.9(–6.0) 5.1 1.0–1.3 1.1

Miettinen 7833,3

& Pippola 30 (4.8–)4.9–7.0(–7.1) 5.6 (4.4–)5.0–7.0 5.5 0.9–1.2 1.0

Kytömäki 3/05 30 (4.4–)4.5–6.4 5.8 (4.1–)4.3–6.1 5.4 0.8–1.3 1.1

Risunen &

Jakobsson 1877 30 3.9–4.8(–4.9) 4.3 (3.3–)3.5–4.4(–4.7) 4.0 1.0–1.2 1.1

T. ramalinae

Haakana 1964 30 (4.5–)4.9–7.4(–7.7) 6.1 (5.7–)5.8–7.8(–7.9) 6.6 0.8–1.1 0.9

Seppälä 1914 30 4.3–6.0(–6.1) 5.5 (4.0–)4.5–6.6(–6.7) 5.8 0.8–1.3 1.0

404 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

n = number of measured structures from given

number of specimens (e.g., 210/7 means 210

structures measured from seven specimens). In

case of size, the entire range is given in paren-

theses. The 90% range excluding the extreme

5% values from both ends is given outside paren-

theses. Where values are identical, parentheses

are omitted. The entire Q range is given without

parentheses.

Biogeographical provinces and collecting

sites are indicated according to Heikinheimo

and Raatikainen (1981). The name “spruce” or

“Picea” refers to Picea abies, “pine” or “Pinus”

to Pinus sylvestris and “birch” or “Betula” to

both Betula pendula and B. pubescens. The dis-

tribution maps were created by Raino Lampinen

(Botanical Museum, Finnish Museum of Natural

History) using the DMAP for Windows software

by Dr. Alan Morton. The maps show the col-

lecting sites in the 10 ¥ 10 km squares of the

Finnish uniform grid coordinate system (KKJ3)

as well as the boundaries of the biogeographi-

cal provinces. If more than one collection of the

same species comes from the same 10 ¥ 10 km

square, only one spot is shown. The illustrations

are based on microscope drawings made with

a drawing tube. Photographs are taken with an

Olympus Camedia C-7070 attached to a binocu-

lar microscope or Sony Cyber-shot DSC-W15.

The nomenclature of Tremella is provided

in the text, along with descriptions of the spe-

cies. The nomenclature of other fungi, includ-

ing lichenized species, follows Index Fungo-

rum (http://www.indexfungorum.org) whilst the

nomenclature of vascular plants follows Tutin

et al. (1964–1980). The authors of the names

are given in these sources and are not generally

repeated here. In citations of the type specimens,

nomenclature is given in its original format, and

an exclamation mark (!) after herbarium acronym

indicates that the type specimen was examined.

The species are arranged in alphabetical order.

Tremella Pers.

Neues Mag. Bot. 1: 111. 1794, nom. cons.

Species of the cosmopolitan genus Tremella

are extremely variable in appearance, including

size, form and colour. Some of the species have

large, gelatinous, foliose, lobate or cerebrifom

basidiomata looking like traditional jelly fungi,

whilst the basidiomata of others are cryptic or

even macroscopically invisible. Lichenicolous

species often grow within galls induced in the

host lichen. Nonetheless, all have characteristics

in common. All the species in the genus appear

to be parasitic: they grow on, in association with,

or in the hymenium of other fungi.

Microscopically the genus is character-

ised by globose to subglobose or ovoid basidia

which become longitudinally or irregularly (e.g.

obliquely or transversely) septate. Basidia are

two- to four-celled or exceptionally one-celled,

and each cell bears an elongated sterigma varia-

ble in length. The hymenium is typically amphig-

enous with abundant probasidia. Spores are glo-

bose to ellipsoid, smooth, thin-walled, hyaline to

slightly coloured and have a distinctive apiculus.

Some spores form secondary spores or germinate

by germ tubes. The yeast phase occurring in the

life cycle originates via budding of the spores or

secondary spores. Hyphae are typically clamped,

but simple-septate hyphae occur in a few spe-

cies. Haustoria are normally present, clamped

and consisting of a haustorial mother cell with

one or more haustorial laments. Hyphidia are

observed in a few species, but usually they are

reduced or absent. Swollen cells are present

in many species, and vesicles are occasionally

observed. Cystidia are absent. Most of the spe-

cies have a conidial stage. Conidia, or in the case

of some lichenicolous species asteroconidia, typ-

ically originate from conidiogenous cells.

Key to the Finnish species of Tremella

1. Basidiomata not macroscopically visible, in the

hymenium of other fungi ............................................ 2.

1. Distinct basidiomata or galls ....................................... 4.

2. In the hymenium of polypores .................. T. polyporina

2. In the hymenium of heterobasidioid Dacrymyces species

..................................................................................... 3.

3. Basidia typically without stalks, conidia present, hyphae

with real clamps ............................................ T. obscura

3. Basidia typically with stalks up to 18 µm long, conidia

absent, hyphae with pseudoclamps ................. T. giraffa

4. On lichens ................................................................... 5.

4. On wood or on non-lichenized fungi ........................ 10.

5. Spores typically ellipsoid (Q* = 1.2), basidiomata pale

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 405

brown to almost black, on Cetrariella delisei or Tucker-

manopsis chlorophylla ............................ T. cetrariicola

5. Spores globose to subglobose, often wider than long (Q*

= 0.9), basidiomata or galls on other lichens .............. 6.

6. Basidia always four-celled, uppermost septum in each

basidium longitudinal or oblique and two lower septa

transverse, basidiomata pale rose to pale brown, on

Ramalina fraxinea ...................................... T. ramalinae

6. Basidia two-celled or very rarely four-celled, all septa

parallel, basidiomata or galls on other lichens ............ 7.

7. Basidia typically with one transverse septum, asteroco-

nidia abundant in some specimens .............................. 8.

7. Basidia longitudinally or obliquely septate ................ 9.

8. Basidiomata or galls olive brown to almost black, on

Phaeophyscia spp. .............................. T. phaeophysciae

8. Basidiomata or galls pale brown to reddish brown, on

Cladonia spp. ............................................. T. cladoniae

9. Basidial cells elongated, variable in size, basidiomata

almost black, on Hypocenomyce scalaris .......................

............................................................ T. hypocenomycis

9. Basidia globose, irregularly ellipsoid, oblong or ovoid,

basidiomata or galls pink to brownish orange, on Hypo-

gymnia physodes ................................... T. hypogymniae

10. Basidiomata mostly more than 1 cm in diam. .......... 11.

10. Basidiomata smaller, only over 1 cm in diam. via coales-

cence ......................................................................... 13.

11. Basidiomata with a rm core, rose-coloured, cerebri-

form ............................................................ T. encephala

11. Basidiomata lacking a rm core, cream, yellow, brown

or almost black, becoming foliose or lobate in maturity

................................................................................... 12.

12. Basidiomata cream, yellow or orange-yellow ................

................................................................. T. mesenterica

12. Basidiomata pale brown to almost black ...... T. foliacea

13. On Juniperus communis ............................... T. karstenii

13. On hosts other than Juniperus communis ................. 14.

14. On Aleurodiscus amorphous on Abies sibirica ...............

.......................................................... T. mycetophiloides

14. On deciduous trees, usually on or together with pyreno-

mycetes (Ascomycota) .............................................. 15.

15. Basidiomata less than 3 mm in diam., basidia long-

stalked ....................................................... T. globispora

15. Basidiomata 3–10 mm in diam., basidia mostly stalkless

................................................................... T. indecorata

Tremella cetrariicola Diederich & Coppins

(Plate 1a, Figs. 1–2)

in Diederich, Bibl. Lichenol. 61: 57. 1996. — Type: UK.

Scotland, Easterness (VC 96), NW of Fort Augustus, Cean-

nacroc Forest, Torgyle Bridge (28/31.13), on Tuckermanop-

sis chlorophylla on Pinus sylvestris, 1.VI.1987 P. Diederich

8864 & B. Coppins [11654] (holotype E; isotypes LG, herb.

Diederich).

Basidiomata cushion-like, discoid or some-

what spherical, 0.2–1.5 mm in diam., typically

with a central depression and, especially in larger

basidiomata, basally constricted, surface smooth,

gelatinous, brown to dark brown or almost black,

rose or pale brown when young, white interior

consisting of lichen thallus.

Spores hyaline, thin-walled, ellipsoid,

(4.2–)5.8–9.6(–10.4) ¥ (3.8–)4.2–7.9(–9.8) µm,

L = 7 µm, W = 5.8 µm, Q = 0.9–1.9, Q* = 1.2

(n = 120/4), apiculus at least partially refractive,

germinating by germ tubes or forming secondary

spores.

Basidia very rarely stalked, 2.4–13.3 ¥ 1.6–

2.4 µm (n = 5/2), two- or infrequently four-

celled, smooth, hyaline, longitudinally, obliquely

or infrequently almost transversely septate, usu-

ally subglobose or ellipsoid, occasionally oblong,

(9.8–)10.2–17.4(–23.1) ¥ (6.5–)6.7–11.9(–12.2)

µm, L = 13.2 µm, W = 9 µm, Q = 1.0–2.9, Q* =

1.5 (n = 58/4), sterigmata up to ca. 32 µm long,

(1.4–)1.8–3.9(–4.1) µm in diam.

Conidia-like cells observed in all specimens,

but they could be secondary spores or haustorial

mother cells. Conidia-like cells smooth, hya-

line, thin-walled, subglobose to oval or ellipsoid,

(2.8–)2.9–6.3(–7.3) ¥ (1.8–)2.3–4(–4.5) µm, L =

4.3 µm, W = 3 µm, Q = 1.0–2.4, Q* = 1.5 (n =

45/4).

All hyphae simple-septate, smooth, hyaline,

(0.8–)1.5–4.1(–5.5) µm in diam. (n = 65/4), thin-

to slightly thick-walled (up to 0.6 µm), occasion-

ally with oil drops. Occasional basidia and haus-

toria with a basal clamp or clamp-like structure.

Haustoria abundant. Haustorial mother

cells smooth, hyaline, subglobose to ellipsoid,

(2.2–)2.4–4.9(–5.5) ¥ (1.9–)2.1–3.7(–4) µm, L

= 3.5 µm, W = 2.8 µm, Q = 0.8–2.4, Q* = 1.3 (n

= 55/4), each mother cell bearing a single haus-

torial lament 0.5–1 µm in diam., up to 12 µm

long, rarely branched.

Swollen cells, vesicles and hyphidia absent.

ecology. In Finland T. cetrariicola is found

on the thalli of Cetrariella delisei and Tuckerma-

nopsis chlorophylla. In addition to these hosts, it

is known to occur on Tuckermanopsis americana

and Tuckermanopsis ciliaris (Diederich 1996).

The former is not reported from Finland and the

latter is extinct (Vitikainen et al. 1997).

The Finnish collections of T. cetrariicola

were made from June to October, and it seems to

prefer spruce-dominated forests.

406 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

DisTribuTion anD abunDance. Though there

are only four collections of T. cetrariicola from

Finland (Fig. 2), we do not consider it a threat-

ened species. Even on the basis of these few

specimens, the distribution of T. cetrariicola

seems to be wide. In addition to the Finnish col-

lections, there is one collection in H from Russia

close to the Finnish border. The host species C.

delisei and T. chlorophylla are widespread and

common in Finland (Vitikainen et al. 1997), and

it supports the idea that the distribution of T.

cetrariicola is wide as well. Tremella cetrarii-

cola has been reported from Sweden, Scotland,

Canary Islands, Canada, the U.S.A. (Diederich

1996), Austria (Triebel 1999), Estonia (Suija

2005) and Russia (Zhurbenko 2007). This is the

rst Finnish record.

noTes. The brown basidiomata with a cen-

tral depression and constricted base as well as

the ellipsoid spores with refractive apiculus

are characteristic of T. cetrariicola. The other

lichenicolous Tremella species have differently

shaped spores.

specimens examineD. — Finland. Etelä-Häme: Riihi-

mäki, on Tuckermanopsis chlorophylla on Picea, 1951 Hir-

venoja (OULU F073608). Pohjois-Savo: Kuopio, Räsälä,

Mustikkasaari, cliff, on Cetrariella delisei, 1909 Linkola

(OULU F073610). Oulun Pohjanmaa: Oulu, Sanginjoki,

Kinnunen, spruce-dominated forest, on T. chlorophylla on

Picea, 2006 Hyvärinen (OULU F073600). Perä-Pohjanmaa:

Tornio, Nivavaara, spruce forest, on T. chlorophylla on twig

of Picea, 2005 Hyvärinen (OULU F073606).

Tremella cladoniae Diederich & M.S. Christ.

in Diederich, Bibl. Lichenol. 61: 65. 1996. — Type: Ger-

Plate 1. — a: Tremella

cetrariicola (arrows),

specimen Hyvärinen

2005, photographed dry.

— b: Tremella encephala,

specimen Pippola 456 with

Stereum sanguinolentum

(arrow), photographed

fresh. — c: Tremella

foliacea, specimen Inarin

Lappi: Inari, Otsamo, on

Betula, 2005 Pippola 343B

(OULU F069518), photo-

graphed fresh. — d: Tre-

mella globispora (arrows),

specimen Söderholm

2752, photographed dry.

— e: Tremella hypoceno-

mycis (arrow), specimen

Norrlin 1867 (holotype),

photographed dry. — f:

Tremella hypogymniae

(arrows), specimen Takala

3, photographed dry.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 407

many. Schwarzwald, Bernau, Spiesshorn, on Cladonia sp.,

1.VIII.1916 G. Lettau s.n. (holotype B 93630; isotype herb.

Diederich).

This is the rst record from Finland. For a detailed

description of the species, see Diederich (1996).

specimen examineD. Finland. Varsinais-Suomi: Halikko,

Vaisakko, rich grass-herb forest, on Cladonia coniocraea on

decaying wood, 2007 Hyvärinen 586 & Syrjänen (H).

Tremella encephala Pers. : Fr. (Plate 1b,

Figs. 2–3)

Syn. Meth. Fung. 2: 623. 1801. — Naematelia encephala

(Pers. : Fr.) Fr., Observ. mycol. 2: 370. 1818; Syst. mycol. 2

(Lundae): 227. 1822.

Tremella encephaliformis Willd., Bot. Mag. (Römer

& Usteri) 2(4): 17. 1788. — Naematelia encephaliformis

(Willd.) Coker, J. Elisha Mitchell Sci. Soc. 35: 137. 1920.

For further synonyms, see Bandoni (1961).

Basidiomata rose- or brownish orange, some-

times slightly greyish, gelatinous, cerebriform, up

to 3 cm in diam., with a white rm core consist-

ing of the hyphae of Stereum sanguinolentum.

Spores hyaline, subglobose, (6–)7.4–11(–12)

¥ (5–)6.2–9.9(–11.6) µm, L = 9.1 µm, W = 8.2

µm, Q = 0.9–1.4, Q* = 1.1 (n = 212/7), apiculus

distinctive, oil drops common, some forming

subglobose to ellipsoid secondary spores or ger-

minating by germ tubes.

Basidia four- or occasionally two-celled,

mostly longitudinally septate, basally clamped,

hyaline, globose or subglobose, (11.8–)12.1–

17.8(–19.8) ¥ (10.5–)12.3–19.6(–21.6) µm, L

= 15.1 µm, W = 16 µm, Q = 0.8–1.2, Q* = 1.0

(n = 114/7), oil drops common, sterigmata up

to ca. 100 µm long, (1.7–)1.9–3.7(–3.9) µm in

diam., frequently apically swollen up to 6.1 µm

in diam.

Fig. 1. Tremella cetrarii-

cola (a–d from Hyvärinen

2005, e–f from Hirven-

oja 1951). — a and e:

Spores, of which one

forms a secondary spore

and one germinates by

a germ tube. — b and

f: Basidia at different

stages of development.

— c: Hyphae with haus-

toria. — d: Conidia-like

cells.

Fig. 2. Distribution of Tremella cetrariicola () and T.

encephala () in Finland. The lines show the bounda-

ries of the biogeographical provinces.

Terminal and subterminal swollen cells abun-

dant close to the substrate and the core, occa-

sional elsewhere. Swollen cells smooth, hyaline,

globose, ellipsoid, citriform, ovoid or oblong,

408 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

(5.1–)5.9–13.6(–14.8) ¥ (4.1–)5.3–9.6(–12) µm,

L = 9.5 µm, W = 7 µm, Q = 1.0–2.4, Q* = 1.4 (n

= 106/7), walls normally up to 0.6 µm, occasion-

ally up to 1.8 µm thick.

Hyphae clamped, smooth, hyaline, (1.2–)1.8–

4.4(–6) µm in diam. (n = 112/7), mostly thin- to

slightly thick-walled (up to 0.6 µm), here and

there very thick-walled (up to 1.8 µm), anas-

tomoses frequent, especially close to the dense

core where the hyphae are intermixed with those

of S. sanguinolentum.

Hyphidia abundant close to the substrate,

to some extent also in hymenium and subhy-

menium. Hyphidia smooth, hyaline, (2.1–)2.4–

5.8(–7.1) µm in diam. (n = 107/7), thin- to

slightly thick-walled (up to 0.4 µm), occasion-

ally thick-walled (up to 1.6 µm). The hyphidia

may be confused with hyphal tips or elongated

swollen cells.

Haustoria abundant close to the core, rare

elsewhere. Haustorial mother cells hyaline,

globose, ellipsoid or oblong, 2.4–5.5(–6.1) ¥

(1.9–)2.1–4.5(–5.5) µm, L = 3.9 µm, W = 3 µm,

Q = 0.9–2.5, Q* = 1.4 (n = 99/7), each bearing

one to four haustorial laments, not more than 1

µm in diam., up to 14 µm long, rarely branched.

Conidial stage and vesicles absent.

ecology. Tremella encephala parasitiz-

ing Stereum sanguinolentum was the rst Tre-

mella species proved to be a parasite (Bandoni

1961). In almost half of the Finnish collections

T. encephala occurs either on (3 specimens stud-

ied) or with (47 specimens studied) basidiomata

of S. sanguinolentum.

The habitats of T. encephala are coniferous

and mixed forest of various ages, and it seems to

be restricted to coniferous wood (Table 2). Even

though T. encephala is occasionally reported to

grow on angiosperms (e.g. Olive 1946a, Bandoni

1961), the single Finnish collection from birch

may be a misidentication. Basidiomata are visi-

ble all year round, but are most commonly found

from September to November.

DisTribuTion anD abunDance. In Finland T.

encephala is commonly found in the boreal zone

where Pinus and Picea occur (Fig. 2). Worldwide

it is recorded in most parts of Europe (Jülich

1984, Krieglsteiner 2000), North America (Ban-

Table 2. Substrates of Tremella encephala, T. foliacea and T. mesenterica.

T. encephala T. foliacea T. mesenterica

Conifers

Picea abies 17 11

Pinus sylvestris 91 8

Unidentiﬁed conifers 1 1

Angiosperms

Alnus spp. 25 30

Betula spp. 1 33 41

Caragana arborescens 1

Corylus avellana 8 2

Fraxinus excelsior 5

Phellodendron amurense 1

Populus tremula 5 7

Prunus padus 1 12

Quercus robur 7 2

Rhamnus frangula 1

Ribes alpinum 1

Salix spp. 3 16

Sambucus racemosa 4

Sorbus aucuparia 1 1

Syringa vulgaris 1

Tilia spp. 2

Ulmus spp. 1

Unidentiﬁed angiosperms 7 23

Unknown substrates 9 26 12

Total 119 137 162

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 409

doni 1961, Krieglsteiner 2000), Japan (Kobayasi

1939, Bandoni 1961, Krieglsteiner 2000), Taiwan

(Chen 1998) and Argentina (Lowy 1971).

noTes. The parasitic relationship gives the

rose-coloured basidiomata of T. encephala a

characteristic feature: the white, rm core is

composed of host hyphae. In addition, basidi-

omata of the host are often present.

selecTeD specimens examineD. — Finland. Uusimaa:

Artjärvi, Hiitelä, Pulikankallio, forest, on fallen Pinus, 2004

Haikonen 23835 (H). Etelä-Häme: Tammela, Mustiala,

on Pinus, Nov. 1881 Karsten (H). Etelä-Savo: Mikkeli,

Vuolinko, on log of Betula, 28 Oct. 1962 Tiensuu (H). Etelä-

Pohjanmaa: Seinäjoki, Hautala, spruce-dominated forest,

on Pinus with Stereum sanguinolentum, 2005 Pippola 456

(OULU F068691). Pohjois-Karjala: Ilomantsi, Ahvensalo,

forest, on twigs of Pinus, 1987 Jäppinen (OULU F044329).

Oulun Pohjanmaa: Oulu, Kaijonharju, Metsänheimo, on

Picea, partially on basidiomata of S. sanguinolentum, 1984

Metsänheimo (OULU F044338). Kittilän Lappi: Kittilä,

Tuuliharju, Moluskoski, on Pinus, 1977 Ohenoja (OULU

F044344).

Tremella foliacea Pers. : Fr. s. lato (Plate

1c, Figs. 4–5)

Observ. mycol. 2: 98. 1799. — Tremella foliacea Pers. : Fr.,

Syst. mycol. 2 (Lundae): 212. 1822. — Gyraria foliacea

(Pers.) Gray, Nat. Arr. Brit. Pl. 1: 594. 1821. — Ulocolla

foliacea (Pers.) Bref., Unters. Gesamtgeb. Mykol. 7: 98.

1888. — Exidia foliacea (Pers.) P. Karst., Bidr. Känn. Finl.

Nat. Folk 48: 449. 1889.

Gyraria ferruginea Gray, Nat. Arr. Brit. Pl. 1: 593.

1821.

Phaeotremella pseudofoliacea Rea, Trans. Br. Mycol.

Fig. 3. Tremella encephala

(a–e from Pippola 456, f

from Jäppinen 1987). — a

and f: Spores, of which

some germinate by a germ

tube or form secondary

spores. — b: Basidia. —

c: Hyphae with haustoria,

of which one is attached to

the host hyphae (arrow).

— d: Swollen cells. — e:

Hyphidia.

410 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

Soc. 3: 377. (1911)1912. — Tremella foliacea var. pseudofo-

liacea (Rea) Kobayasi, Sci. Rep. Tokyo Bunrika Daigaku B

4: 23. 1939. — Type (Roberts 1999): UK. England, Somerset,

Staple Park, 20 Sep. 1911 C. Rea (lectotype K(M) 56574).

— Synonymized by Donk (1966) and Roberts (1999).

Tremella mbriata Pers. : Fr., Observ. mycol. 2: 97.

1799. — Tremella mbriata Pers. : Fr., Syst. mycol. 2

(Lundae): 212. 1822. — Tremella foliacea var. mbriata

(Pers.) S. Lundell, Fungi Exs. Suecici Praes. Upsal. 19-20:

16 (no. 940). 1941.

Tremella nigrescens Fr., Summa veg. Scand.: 341. 1849.

— Exidia nigrescens (Fr.) P. Karst., Bidr. Känn. Finl. Nat.

Folk 48: 449. 1889. — Synonymized by Roberts (1999).

Tremella succinea Pers., Mycol. Eur. 1: 101. 1822.

— Tremella foliacea var. succinea (Pers.) Neuhoff, Z. Pilzk.

10: 73. 1931.

Tremella crispa Lloyd, Mycol. Writ. 7: 1152. 1922.

— Type: Australia. Tasmania, Hobart, undated L. Rodway

(herb. Lloyd 4004) (holotype BPI ). — Synonymized by

Bandoni (1958).

? Tremella frondosa Fr., Syst. mycol. 2 (Lundae): 212.

1822. — Proposals for synonymy made e.g. by Coker (1920)

and Looney (1933), but T. frondosa is also suggested to be

synonymous with T. aurantia Schwein. : Fr. (Roberts 1995).

Basidiomata gelatinous, pale brown to almost

black, becoming foliate or lobate in maturity,

from one to over 10 cm in diam.

Spores hyaline to pale brown, globose to ellip-

soid, (6–)6.5–10(–11.4) ¥ (4.5–)5.9–9.4(–9.8)

µm, L = 8.2 µm, W = 7.3 µm, Q = 0.9–1.5, Q*

= 1.1 (n = 210/7), apiculus distinctive, oil drops

common, some forming secondary spores or ger-

minating by germ tubes.

Basidia four- or occasionally two-celled, lon-

gitudinally of obliquely septate, basally clamped,

hyaline to pale brown, globose to subglobose

or ovoid, (9.4–)11–19.2(–20.2) ¥ (9.4–)10–

15.9(–19.8) µm, L = 14.1 µm, W = 13.4 µm,

Q = 0.7–1.9, Q* = 1.1 (n = 132/7), oil drops

common, infrequently new basidia originating

Plate 2. — a: Tremella

indecorata (right arrow),

specimen Pippo la 80

on Diatrype bullata (left

arrow), photographed dry.

— b: Tremella karstenii,

specimen Inarin Lappi:

Utsjoki, Paistunturit, on

Juniperus communis,

2005 Pippola 398 (OULU

F069353), photographed

fresh. — c: Tremella

mesenterica, specimen

Inarin Lappi: Utsjoki,

Paistunturit, on Betula,

2005 Knuuttila (OULU

F069526), photographed

fresh. — d: Tremella

mycetophiloides (arrows),

specimen Kotiranta

11860, photographed dry.

— e: Tremella phaeophy-

sciae (arrow), specimen

Hausen 1927 (OULU

F073618), photographed

dry. — f: Tremella rama-

linae (arrow), specimen

Haakana 1964, photo-

graphed dry.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 411

from existing basidia, sterigmata up to ca. 75 µm

long, (1.6–)1.8–3.8(–3.9) µm in diam., often api-

cally swollen up to 5.8 µm in diam.

Conidia commonly, but not always, observed

close to the substrate and occasionally elsewhere,

originating from swollen or conidiogenous cells.

Conidia hyaline, smooth, thin-walled, glo-

bose, ellipsoid or oblong, (1.9–)2.4–5.4(–5.9)

¥ (1.1–)1.6–3.8(–4.2) µm, L = 3.9 µm, W = 2.3

µm, Q = 1.0–3.1, Q* = 1.8 (n = 120/4).

Terminal and subterminal swollen cells

observed especially close to the substrate. Swol-

len cells smooth, hyaline, pale brown or brown,

variable in size and form, globose, ellipsoid,

oblong or cylindrical, (5.1–)6.5–39.1(–53.3) ¥

(2.3–)5.7–31(–51.1) µm, L = 14.7 µm, W = 11.1

µm, Q = 0.9–2.6, Q* = 1.4 (n = 147/7), thin- to

very thick-walled (up to 1.6 µm), occasionally

extremely thick-walled (up to ca. 4 µm).

Hyphae clamped, smooth or gelatinous, hya-

line to pale brown, (1.5–)1.9–6.3(–11.8) µm in

diam. (n = 117/7), thin- to very thick-walled (up

to 1.8 µm).

Haustoria few or abundant, occurring espe-

cially close to the substrate, originating from

hyphae or swollen cells. Haustorial mother cells

basally clamped, hyaline or pale brown, subglo-

bose, ellipsoid or oblong, (3.4–)3.8–6.1(–6.9) ¥

(2.1–)2.4–4.1(–5.5) µm, L = 4.6 µm, W = 3.4 µm,

Q = 1.4–2.9, Q* = 1.4 (n = 93/7), each bearing one

or a few haustorial laments, 0.5–1.5 µm in diam.,

up to ca. 60 µm long, branched or unbranched.

Fig. 4. Tremella foliacea

(a–d from Malmström

39, e from Pippola 376, f

from Koskela 1970). — a:

Spores, of which two ger-

minate by germ tubes. —

b: Basidia. — c: Hyphae

with haustoria and a few

swollen cells. — d, e and

f: Various swollen cells,

of which some produce

conidia.

412 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

Vesicles and hyphidia absent.

ecology. Tremella foliacea is presumably a

parasite of Stereum species (Roberts 1999, Tor-

kelsen 2005). In Finland it is noted together with

Stereum hirsutum (2 specimens studied), Stereum

rugosum (4 specimens studied) and Stereum sub-

tomentosum (1 specimen studied). Tremella folia-

cea is clearly more common on deciduous wood,

but occurs occasionally on conifers (Table 2). Its

basidiomata exist all year round. Most commonly

they are found from August to October, and they

are edible (Boa 2004). Tremella foliacea seems to

prefer herb-rich forests, but it occurs frequently

also in pastures, gardens, coniferous forests, as

well as in peatlands and virgin forests.

DisTribuTion anD abunDance. Tremella folia-

cea is a common species found almost through-

out Finland (Fig. 5). Worldwide it is known from

North (Coker 1920, Martin 1952), Central (Lowy

1971) and South America (Roberts 1999), Aus-

tralia (Roberts 1999), Asia (Kobayasi 1939, Teng

1996, Chen 1998), Africa (Wojewoda 1981) and

Europe (e.g. Wojewoda 1981, Jülich 1984, Tor-

kelsen 1997, Roberts 1999).

noTes. Brown to almost black, foliose basidi-

omata are specic to T. foliacea. However, it is

a very variable species in many respects such

as substrate, colour, presence of conidia and

spore size. For instance, depending on author

the conidia are either absent (e.g. Neuhoff 1931,

Martin 1952, Chen 1998) or present (e.g. Koba-

yasi 1939, Lowy 1971, Wojewoda 1981, Rob-

erts 1999). Because of the huge variability, T.

foliacea has been described several times with

numerous variations. Now most of them are

regarded as synonyms, but some might be good

taxa. For instance, it is not sure if T. foliacea and

T. frondosa are conspecic. If they are distinct,

T. frondosa is distinguished from T. foliacea by

its yellow to pale brown colour and presence of

conidia (Chen 1998), or it may be synonymous

with T. aurantia (Roberts 1995). Alternatively,

the paler colour may be a result of exposure to

liquid or rain (Kobayasi 1939).

Since the concept of the species is not well

established, we prefer to use T. foliacea s. lato.

As noted by Chen (1998), a good specimen

should be selected as a neotype for T. foliacea

from Europe where it was originally described.

Such a neotypication could help the study of

this complex.

selecTeD specimens examineD. — Finland. Åland: Lem-

land, Marsö, on fallen Quercus robur, 1948 Stenlid 668

(UPS). Uusimaa: Siuntio, centre, on Alnus incana, 23.X.1966

Oittinen (H). Etelä-Savo: Kangasniemi, Luusniemi, Siko-

salo, on Betula, 1990 Malmström 39 (H). Pohjois-Karjala:

Kitee, Kiteenjärvi, Potoskanlahti, on stump of Betula, 1966

Kankainen (TUR 28148). Keski-Pohjanmaa: Haapavesi,

birch forest, on Picea, 1970 Koskela (OULU F044347).

Koillismaa: Taivalkoski, Kylmälä, Moskavaara, on fallen

Picea, 1972 Ohenoja & Ohenoja (OULU F044355). Inarin

Lappi: Utsjoki, Kistuskaidi, mountain birch forest, on fallen

Salix, 2005 Pippola 376 (OULU F069358).

Tremella giraffa Chee J. Chen (Figs. 6–7)

Bibl. Mycol. 174: 173. 1998. — Type: Germany. Tübingen,

Fig. 5. Distribution of Tremella foliacea () in Finland.

The lines show the boundaries of the biogeographical

provinces.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 413

Schönbuch, in basidiocarps of Dacrymyces stillatus grow-

ing on branches of Larix decidua, 29.VII.1997 R. Kirschner

(holotype TUB; isotype TAI).

Basidiomata not macroscopically visible, in

the host hymenium.

Spores smooth, hyaline, thin-walled, glo-

bose to ellipsoid, (5.7–)6.3–8.8(–9.9) ¥ (5–)5.9–

8(–9.8) µm, L = 7.6 µm, W = 6.9 µm, Q = 0.9–

1.4, Q* = 1.1 (n = 61/2), apiculus distinctive, oil

drops common, some germinating by germ tubes

or forming secondary spores.

Basidia usually stalked, (2–)2.1–17(–18) ¥

(1.1–)1.8–3.8 µm (n = 31/2), two-celled, smooth,

hyaline, thin-walled, longitudinally or occasion-

ally obliquely or transversely septate, globose

to ellipsoid, (9.4–)9.8–14.9(–15.1) ¥ (8.4–)8.8–

13.1(–14) µm, L = 12.5 µm, W = 11.3 µm, Q =

0.9–1.4, Q* = 1.1 (n = 33/2), sterigmata up to

ca. 50 µm long, (2.1–)2.2–4(–4.8) µm in diam.,

often apically swollen up to 5.9 µm in diam.

Hyphae smooth, hyaline, (1.7–)1.8–4.2(–4.3)

µm in diam. (n = 32/2), thin-walled (under 0.2

µm), oil drops common. Clamp-like structures,

so-called pseudoclamps, near the septa observed

instead of real clamps.

Haustoria abundant, originating from hyphae,

basidia or stalks of basidia. Haustorial mother cells

smooth, hyaline, thin-walled, globose to ellipsoid,

(2.3–)2.9–4(–4.2) ¥ 2.1–3.2 µm, L = 3.4 µm, W

= 2.7 µm, Q = 1.0–1.9, Q* = 1.3 (n = 30/2), each

mother cell bearing one to three haustorial la-

ments 0.5–1 µm in diam., up to 21 µm long.

Fig. 6. Tremella giraffa (a–

d from Alanko 132586).

— a: Spores, of which one

is germinating by a germ

tube. — b: Basidia. — c:

Haustoria and new basidia

originating from basidia.

— d: Pseudoclamped

hyphae with haustoria and

a probasidium.

Fig. 7. Distribution of Tremella giraffa (), T. globispora

() and T. hypocenomycis (+) in Finland. The lines

show the boundaries of the biogeographical provinces.

414 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

Conidial stage, swollen cells, vesicles and

hyphidia absent.

ecology. Tremella giraffa is parasitic in the

hymenium of Dacrymyces species. In Finland it

is found in the hymenium of Dacrymyces minor.

It is also known to parasitize D. stillatus (Chen

1998) and D. capitatus (Van de Put 2000).

DisTribuTion anD abunDance. This is the rst

record of T. giraffa from Finland. So far, it

is known only from two localities (Fig. 7).

Even though Dacrymyces species are deciently

known in Finland, they seem to be common and

widespread. Tremella giraffa may be common

and widespread as well. However, further stud-

ies are needed to understand its distribution

and evaluate if it is a threatened, rare or abun-

dant species. In addition to the type locality in

Germany (Chen 1998), T. giraffa is found only

in England, Hungary, Italy (Roberts 2007) and

Belgium (Van de Put 2000), and it is deciently

known worldwide.

noTes. In addition to T. giraffa, there are

three other Tremella species parasitizing Dacry-

myces species: T. obscura, T. occultifuroidea and

T. penetrans. Tremella giraffa is distinguished

from the others especially by its pseudoclamps

which are reminiscent of real clamps, but are not

fully developed. Moreover, most of its basidia

are stalked, haustoria as well as new basidia

originate occasionally directly from basidia, and

conidia are typically, but not necessarily (see

Van de Put 2000), absent.

specimens examineD. — Finland. Uusimaa: Helsinki,

Pitäjänmäki, Marttila, in Dacrymyces minor on Sorbus

aucuparia, 2007 Alanko 132586 (H). Etelä-Häme: Hämeen-

linna, railway station NW, in D. minor on Salix caprea, 2007

Alanko 132671 (H).

Tremella globispora D.A. Reid (Plate 1d,

Figs. 7–8)

Trans. Br. Mycol. Soc. 55: 414. 1970, as ‘globospora’.

— Type: UK. W. Sussex, West Dean Wood, West Dean, on

Rubus fruticosus agg., in association with Diaporthe eres,

21.XII.1968 D. A. Reid (holotype K).

Tremella candida var. globispora (D.A. Reid) Krieglst.,

Beitr. Kenntn. Pilze Mitteleur. 12: 39. 1999.

Tremella tubercularia Berk. sensu Bourdot and Galzin

(1927), Neuhoff (1931), etc.

Basidiomata cushion-like and usually some-

what cerebriform, 1–3 mm in diam., larger

only via coalescence, black when dry, greyish

to blackish brown and slightly hyaline when

soaked

Spores hyaline to brown, thin-walled, glo-

bose to subglobose, often wider than long,

(5.4–)5.9–9.6(–10.2) ¥ (5.4–)5.9–10.2(–12) µm,

L = 7.6 µm, W = 8.1 µm, Q = 0.8–1.2, Q* = 1.0

(n = 90/3), apiculus distinctive.

Fig. 8. Tremella globis-

pora (a–c from Söderholm

2752, d from Karsten

1862). — a: Spores. — b:

Basidia at different stages

of development. — c:

Hyphae with a swollen cell

and haustoria. — d: Wide

and thick-walled hyphae

close to the substrate.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 415

Basidia usually stalked, (2.4–)3.7–

24.7(–27.8) ¥ (1.9–)2–4.5(–5.4) µm (n = 50/3),

basally clamped, two- or four-celled, longitu-

dinally, obliquely or occasionally transversely

septate, hyaline, pale brown or brown, some-

times thick- or very thick-walled (up to 2 µm),

subglobose to ellipsoid, (12.3–)13.7–21.3(–22)

¥ (11.4–)12–20(–20.8) µm. L = 17 µm, W = 15.3

µm, Q = 0.8–1.7, Q* = 1.1 (n = 51/3), sterig-

mata up to 63 µm long, (1.4–)1.8–4(–4.5) µm in

diam., often collapsed.

A few swollen cells observed in subhy-

menium close to the substrate, pale brown to

brown, globose, ovoid or ellipsoid, (5.5–)5.8–

10(–11.4) ¥ (4–)4.3–7.9(–8.4) µm, L = 7.8 µm,

W = 6.2 µm, Q = 0.9–1.9, Q* = 1.3 (n = 45/3),

thin- to slightly thick-walled (up to 0.4 µm).

Hyphae clamped, smooth, hyaline, pale

brown or brown, (1.4–)1.7–6.1(–8.1) µm in

diam. (n = 65/3), thin- to very thick-walled

(up to 2 µm). Wide and thick-walled hyphae

common especially close to the substrate.

Haustorial mother cells hyaline to pale

brown, globose to ellipsoid or oblong, (1.9–)2.2–

5.7(–6.1) ¥ (1.8–)2–3.3(–3.5) µm, L = 3.5 µm, W

= 2.3 µm, Q = 0.8–2.6, Q* = 1.5 (n = 45/3), each

mother cell bearing one or a few haustorial la-

ments up to 1 µm in diam. and 18 µm long.

Conidia, vesicles and hyphidia absent.

ecology. Tremella globispora occurs in

association with pyrenomycetes (Ascomycota).

All Finnish collections come from Salix spp.,

and one of them was found on Allantoporthe

tessella. (syn. Diaporthe tessella). According to

the original description (Reid 1970), T. globi-

spora is constantly associated with the perithecia

of Diaporthe spp. though perithecia may be

indistinct. In addition to Diaporthe species, pos-

sible host species are known in the genera Valsa

(Brough 1974) and Eutypella (Wojewoda 1981).

As the hosts infest various deciduous trees, T.

globispora may be found not on only Salix spp.

but also on other deciduous trees.

Finnish specimens were collected either in

January or in October. Thus, basidiomata of T.

globispora may be visible all year round. At

least this is the case in British Columbia (Brough

1974).

DisTribuTion anD abunDance. So far, ndings

are too few (Fig. 7) to reveal the actual distribu-

tion and abundance of T. globispora in Finland

and assist in evaluating if the species is threat-

ened. In general T. globispora seems to be fairly

rare, but widespread. It is found in most parts of

Europe (e.g. Bourdot & Galzin 1927, Neuhoff

1931, Reid 1970, Wojewoda 1981, Jülich 1984,

Torkelsen 1997), North America (Brough 1974),

Colombia (Lowy 1971) and China (Zhuang

2005). This is the rst Finnish record.

noTes. The small basidiomata, only a few

millimetres across, which are associated with

perithecia of pyrenomycetous fungi, characterize

T. globispora. It is distinguished microscopically

from T. indecorata, another Finnish Tremella

species associated with pyrenomycetes, by its

typically stalked basidia. However, the differ-

ence is not necessarily well established (Legon

et al. 2005, P. Roberts personal comm.).

Conidia are absent in Finnish T. globispora

specimens, but according to Chen (1998) they

do exist.

Sebacina globospora Whelden is not a syno-

nym of T. globispora as e.g. Reid (1970) sus-

pected. Both are associated with pyrenomyc-

etous Diaporthe spp., but S. globospora lacks

clamp-connections (Whelden 1935). Sebacina

globospora is currently known as Tremella

diaporthicola Ginns & M.N.L. Lefebvre (Ginns

& Lefebvre 1993).

specimens examineD. — Finland. Etelä-Häme: Lem-

päälä, Hulikankulma, on old Allantoporthe tessella on dead

Salix twigs, 1998 Söderholm 2752 (TUR 122476). Tam-

mela, Mustiala, on twigs of Salix phylicifolia, 7.X.1867

Karsten (H). Keski-Pohjanmaa: Pietarsaari, on twigs of Salix,

24.X.1862 Karsten (H, S F44064).

Tremella hypocenomycis Diederich (Plate

1e, Figs. 7 and 9)

Bibl. Lichenol. 61: 87. 1996. — Type: Finland. Perä-Pohjan-

maa, Pello, Turtola, on Hypocenomyce scalaris, 1867 J.P.

Norrlin (holotype H!; isotype herb. Diederich).

Basidiomata gelatinous, somewhat irregular,

grainy or cerebriform, 0.2–2 mm in diam., black-

ish brown to black when dry, pale brown to

blackish brown when soaked.

Spores smooth, thin-walled, hyaline to slightly

pale brown, globose or subglobose, (4.6–)5.4–

7.6(–7.8) ¥ (4–)5.3–7.6(–8.1) µm, L = 6.4 µm,

416 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

W = 6.2 µm, Q = 0.9–1.2, Q* = 1.0 (n = 20/2),

apiculus remarkably broad.

Basidia two-celled, cells elongated and very

variable in size, smooth, hyaline to pale brown,

longitudinally or exceptionally obliquely septate,

at the septum (7.4–)8.4–17.6(–18.5) ¥ (9.6–)9.8–

21.1(–23.7) µm, L = 12.4 µm, W = 13.7 µm, Q =

0.5–1.5, Q* = 0.9 (n = 33/2), above the septum

individual cells 2.4–7.4(–10.2) µm in diam., up

to 25.5 µm long, sterigmata absent, possibly

degenerated.

All hyphae simple-septate. Two kinds of

hyphae observed: so-called context hyphae hya-

line to pale brown, smooth, 1.1–2.7 µm in diam.

(n = 34/2), thin-walled, so-called fertile hyphae

usually pale brown to brown, smooth, (2.3–)2.7–

8.3(–8.6) µm in diam. (n = 50/2), thick-walled

(up to 1.4 µm), individual cells short, sometimes

almost globose.

Haustoria extremely abundant, originating

mostly from fertile hyphae. Haustorial mother

cells hyaline to brown, globose or subglobose,

(1.8–)2–4.4(–5.3) ¥ (2–)2.1–5.1(–5.6) µm, L =

3.1 µm, W = 3.3 µm, Q = 0.6–1.6, Q* = 1.0 (n =

36/2), sometimes with walls slightly thickened

(up to 0.4 µm), each mother cell bearing one

or occasionally a few haustorial laments up

to 1 µm in diam. and 10.5 µm long, sometimes

slightly branched.

Conidial stage, swollen cells, vesicles and

hyphidia absent.

ecology. Tremella hypocenomycis occurs on

the thallus of Hypocenomyce scalaris.

DisTribuTion anD abunDance. Until now, T.

hypocenomycis was only known from the type

locality. The specimen from Alavus (western Fin-

land) is thus only the second record in the world

(Fig. 7). Both specimens are old, and one might

suggest the species is extinct. However, in the

light of the distribution and abundance of its host

(Vitikainen et al. 1997), this possibility seems to

be unlikely. New specimens of T. hypocenomycis

should be found to better understand its actual

distribution, abundance and potential threats. At

the moment the Red List status of T. hypoceno-

mycis is impossible to evaluate.

noTes. The unusual basidia with elongated

cells at maturity characterize T. hypocenomy-

cis. Another lichenicolous Tremella species with

similar basidia, T. christiansenii Diederich, has

larger spores, viz. 9–12 ¥ 8.5–10.5 µm (Died-

erich 1996).

specimens examineD. — Finland. Etelä-Pohjanmaa:

Alavus, Niinimaa, Uusi-Erkkilä, on Hypocenomyce scalaris

on charred trunk, 1947 Leppälä (OULU F073619). Perä-

Pohjanmaa: Pello, Turtola, on H. scalaris, 1867 Norrlin

(holotype H).

Tremella hypogymniae Diederich & M.S.

Christ. (Plate 1f, Figs. 10–11)

in Diederich, Bibl. Lichenol. 61: 90. 1996. — Type: France.

Pyrénées-Atlantiques, au sud de Arette, un peu en aval du

Chalet d’Oumarre, on Hypogymnia physodes, 29.VII.1990 P.

Diederich 9145 (holotype LG; isotype herb. Diederich).

Inducing galls or basidiomata on the host

thallus. Galls frequent, pale yellow, pinkish,

rose or brownish orange, many at least partially

blackened, somewhat spherical, 0.1–2.5 mm in

diam., with a thin, gelatinous layer on the surface

and a white interior consisting of lichen thal-

Fig. 9. Tremella hypocen-

omycis (a–d from Nor-

rlin 1867). — a: Spores.

— b: Basidia at different

stages of development.

— c: Context hyphae with

haustoria. — d: Fertile

hyphae with haustoria.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 417

lus. Basidiomata occasional, orange or brownish

orange, slightly cerebriform, gelatinous.

Spores smooth, hyaline, thin- to relatively

thin-walled, globose to subglobose, mostly wider

than long, (4.1–)5.1–7.9(–8.3) ¥ (4.9–)5.9–

9.3(–10) µm, L = 6.4 µm, W = 7.4 µm, Q =

0.6–1.3, Q* = 0.9 (n = 179/7), distinct apiculus

frequently refractive, spores germinating by

germ tubes.

Basidia extremely rarely short-stalked, 3.6–

4.3 ¥ 3.6–3.7 µm (n = 2/2), two- or exceptionally

four-celled, cells sometimes variable in size,

hyaline, longitudinally, obliquely or occasionally

transversely septate, globose, irregularly ellip-

soid, oblong or ovoid, (9.8–)11.6–19.6(–23.5)

¥ (7.9–)9.8–14.1(–15.9) µm, L = 14.8 µm, W =

12.1 µm, Q = 0.7–2.3, Q* = 1.2 (n = 107/7), oil

drops common, sterigmata up to 34 µm long,

(1.8–)2–5.4(–5.7) µm in diam.

In some specimens hyphae clamped, in others

mostly or completely simple-septate. Hyphae

hyaline, smooth, (1.8–)2–5.8(–7.8) µm in diam.

(n = 114/7), thin- to very thick-walled (up to 2

µm), mostly with slightly thickened walls (0.2–

0.6 µm), sometimes with oil drops. Hyphae,

especially in the inner parts of the galls, inter-

mixed with hyphae and other structures of the

host lichen.

Haustoria normally few, occasionally either

absent or abundant. Haustorial mother cells hya-

line, globose to ellipsoid, (3.7–)3.8–6.1(–7.4) ¥

Fig. 10. Tremella hypo-

gymniae (a–c from Linkola

1930, d–e from Takala

73). — a: Spores, of which

one is germinating by a

germ tube. — b and d:

Basidia at different stages

of development. — c:

Clamped hyphae with

haustoria. — e: Simple-

septate hyphae.

Fig. 11. Distribution of Tremella hypogymniae () in

Finland. The lines show the boundaries of the biogeo-

graphical provinces.

418 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

(2.9–)3.5–4.5(–5.9) µm, L = 5 µm, W = 4 µm,

Q = 1.0–1.9, Q* = 1.3 (n = 51/6), each mother

cell bearing one haustorial lament 0.5–2 µm

in diam., up to 23 µm long, rarely slightly

branched.

Conidial stage, swollen cells, vesicles and

hyphidia absent.

ecology. Galls and basidiomata of T. hypo-

gymniae are only found on the thallus of Hypo-

gymnia physodes, an extremely common lichen

species occuring on various substrates, espe-

cially on trunks and twigs, but also on iron scrap,

stones, etc. Basidiomata of T. hypogymniae are

found in Finland almost all year round, most fre-

quently from June to August.

DisTribuTion anD abunDance. Tremella hypo-

gymniae is common in the whole country (Fig.

11), and it is widespread in Europe (Diederich

1996). It is also reported from Canada (Diederich

1996, 2003) and the U.S.A. (Diederich 2003)

including Alaska (Geiser et al. 1998), but seems

to be rare in North America (Diederich 2003).

noTes. Pale rose to brownish galls on the

host thallus characterize T. hypogymniae. The

predominantly two-celled basidia of T. hypogym-

niae are smaller than those of Tremella lobar-

iacearum Diederich & M.S. Christ., Tremella

phaeographidis Diederich, Coppins & Bandoni

and other lichenicolous Tremella species which

are microscopically similar (Diederich 1996).

In spite of the variability in hyphal septa, all

the Finnish specimens clearly belong to a single

species. Spore sizes as well as other micro-

scopical structures are identical to each other.

Basidiomata of various ages possibly differ in

their septa. Simple-septate hyphae were com-

monly observed especially in young basidiomata

with plenty of probasidia and only a few mature

basidia and spores.

selecTeD specimens examineD. — Finland. Varsinais-

Suomi: Salo, Uskela, Pahkavuori, on Hypogymnia physodes

on Picea, 26.VI.1919 Häyrén (H). Uusimaa: Orimattila, on H.

physodes on Salix caprea, 1965 Takala 73 (H). Etelä-Karjala:

Ylämaa, Kavenoja, on H. physodes on Betula, 1965 Takala

82 (H). Pohjois-Savo: Pieksämäki, Jäppilä, on H. physodes

on Betula, 1964 Takala 3 (H). Perä-Pohjanmaa: Ylitornio,

Aavasaksa, on H. physodes on Pinus, 21.VII.1936 Räsänen

(H). Koillismaa: Kuusamo, Oulankajoki, Kiutaköngäs, on

H. physodes on Picea, 11.VIII.1930 Linkola (H). Enontekiön

Lappi: Enontekiö, Hetta, Jyppyrävaara, on H. physodes on

Betula, 1965 Takala 103 (H).

Tremella indecorata Sommerf. (Plate 2a,

Figs. 12–13)

Suppl. . lapp.: 306. 1826. — Exidia indecorata (Sommerf.)

P. Karst., Rev. Mycol. (Toulouse) 12: 126. 1890. — Type:

Norway. Nordland, Saltdal, on dead branches of Salix phyl-

icifolia, III.1824 S. C. Sommerfelt (holotype O 72784!).

Basidiomata translucent pale brown to brown,

cerebriform or cushion-like, 3–10 mm in diam.,

occasionally up to 20 mm via coalescence, upon

drying turning to thin, dark brown or black lms.

Spores smooth, hyaline, pale brown or brown,

thin-walled, globose to subglobose, often wider

than long, (4.9–)5.9–9.4(–9.9) ¥ (5.7–)5.9–

9.8(–11.2) µm, L = 7.4 µm, W = 7.7 µm, Q =

0.8–1.3, Q* = 1.0 (n = 150/5), distinct apiculus

often refractive, germinating by germ tubes or

forming globose to ellipsoid secondary spores.

Basidia rarely stalked, 2–12 ¥ 2.2–4 µm

(n = 13/4), two- or four-celled, longitudinally

or obliquely septate, basally clamped, smooth,

hyaline to brown, globose, ellipsoid or citri-

form, sometimes wider than long, (11.8–)12.2–

19.6(–27.1) ¥ (11.9–)13.1–20(–22) µm, L = 16.1

µm, W = 16.2 µm, Q = 0.8–1.7, Q* = 1.0 (n

= 89/5), sterigmata up to ca. 100 µm long,

(1.5–)1.7–4(–5.7) µm in diam., often collapsed,

apical protuberances and thick walls (up to 1.8

µm) observed especially in probasidia, oil drops

common.

Conidial stage absent in most specimens.

Conidia hyaline, oblong or cylindrical, (2.4–)2.6–

4.2 ¥ 0.8–1.6 µm, L = 3.2 µm, W = 1 µm, Q =

1.7–5.3, Q* = 3.5 (n = 30/1). Conidia originat-

ing from conidiogenous cells which are hya-

line to brown, subglobose, ellipsoid or oblong,

(1.9–)2.4–4.4(–5.9) ¥ (1.7–)1.8–3.4 µm, L =

3.6 µm, W = 2.7 µm, Q = 1.1–2.0, Q* = 1.3 (n =

30/1), thin-walled, small oil drops common.

Terminal and subterminal swollen cells

observed in hymenium and subhymenium. Swol-

len cells hyaline to brown, smooth, usually thick-

walled (up to 2 µm), globose, ellipsoid, ovoid,

oblong or citriform, (5.9–)6.5–15.9(–17.8) ¥

(4.3–)5–9.7(–9.9) µm, L = 10.5 µm, W = 7.1

µm, Q = 1.0–2.8, Q* = 1.5 (n = 77/5). Hyphae or

basidia originate occasionally from swollen cells

which may be mistaken for probasidia.

Hyphae clamped, smooth or gelatinous, hya-

line to brown, (1.4–)1.6–5.9(–7.6) µm in diam.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 419

(n = 103/5), thin- to very thick-walled (up to 2

µm).

Haustoria few or absent. Haustorial mother

cells hyaline to pale brown, globose to ellipsoid,

2.5–5.7 ¥ 2.2–4.3 µm, L = 4 µm, W = 3 µm, Q

= 0.9–1.7, Q* = 1.3 (n = 19/2), each mother cell

bearing one haustorial lament, mostly under 1

µm in diam., up to 12 µm long, rarely branched.

Vesicles and hyphidia absent.

ecology. In Finland T. indecorata is found

on deciduous wood, often on Diatrype bullata.

A connection with at least remnants of a pyreno-

mycete (dead perithecia or traces of mycelium)

should always be present (Torkelsen 1968). In

addition to Diatrype species, possible host spe-

cies are known in the genus Eutypa (Torkelsen

1997). All Finnish specimens were collected

either in October or November.

DisTribuTion anD abunDance. Even though

there are only ve collections from two biogeo-

graphical provinces (Fig. 13), the host species

are so common and widespread in Finland (Diss-

ing 2000) that we cannot presume T. indecorata

to be an especially rare species. Worldwide T.

indecorata seems to be restricted in Europe

where it is reported e.g. from the Nordic coun-

tries (Torkelsen 1997), Estonia (Jülich 1984),

Poland (Wojewoda 1981) and Spain (Dueñas

1997).

noTes. Tremella indecorata is one of the two

Finnish Tremella species occurring together with

pyrenomycetes and forming comparatively small

basidiomata. Unlike T. globispora, T. indecorata

mostly has stalkless basidia. In addition, the

basidiomata of T. indecorata seem to be slightly

larger than those of T. globispora.

There are also other tiny Tremella species

associated with pyrenomycetous fungi, such as

T. moriformis Berk. and T. subanomala Coker,

which resemble T. indecorata. On the basis

Fig. 12. Tremella indeco-

rata (a–c from Pippola

80, d–e from Söderholm

2686, f from Haikonen

1866). — a: Spores and

secondary spores. — b

and d: Basidia at differ-

ent stages of develop-

ment. — c: Hyphae with

a few swollen cells. — e:

Hyphae with haustoria. —

f: Conidia originating from

conidiogenous cells.

420 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

needed to reveal the actual relationships and the

systematic position of the species in the Indeco-

rata group.

specimens examineD. — Finland. Etelä-Häme: Asikkala,

Kolunkulma, herb-rich forest, on Corylus avellana, 1981

Haikonen 1866 (H). Lempäälä, Sarvikaslahti, on old Dia-

trype bullata on twigs of Salix, 1996 Söderholm 2686

(OULU F034569). Tammela, Mustiala, on a twig of Betula,

2.XI.1865 Karsten (H). Oulun Pohjanmaa: Oulu, Hietasaari,

Johteensalmi, willow thicket, on Diatrype bullata on dead

twig of Salix caprea, 2004 Pippola 80 (OULU F074015).

Oulu, Hietasaari, Johteensalmi, herb-rich forest, on living S.

caprea, 2004 Pippola 115 (OULU F074016).

Tremella karstenii Hauerslev (Plate 2b,

Figs. 13–14)

Mycotaxon 72: 480. 1999. — Tremella juniperina P. Karst.,

Fungi Fenn. exs. 812. 1869, nomen illeg. (non Tremella juni-

perina L.). — Exidia juniperina P. Karst., Bidr. Känn. Finl.

Nat. Folk 48: 452. 1889. — Type: Finland. Etelä-Häme, Tam-

mela, Mustiala, på en [on Juniperus communis], 8.IV.1869 P.

A. Karsten (lectotype H!; isolectotypes H!).

Basidiomata hyaline to yellowish, smooth

or slightly papillose, gelatinous, cushion-like or

spherical, sometimes partially resupinate, 1–5

mm in diam., drying to thin, tough, almost invis-

ible lms.

Spores hyaline, smooth, thin-walled, glo-

bose, often wider than long, (2.5–)3.4–6(–7.6)

¥ (3–)3.5–6.2(–7.5) µm, L = 4.5 µm, W = 4.9

µm, Q = 0.7–1.4, Q* = 0.9 (n = 210/7), apicu-

lus distinctive, occasionally refractive, oil drops

common, some forming subglobose to ellipsoid

secondary spores.

Basidia mostly four-celled, rarely two-celled,

longitudinally or occasionally obliquely or

transversely septate, basally clamped, hyaline,

smooth, globose to ellipsoid, sometimes wider

than long, (7.8–)8.4–12.5(–15.7) ¥ (7.7–)7.9–

13.3(–15.3) µm, L = 10.5 µm, W = 10.4 µm, Q

= 0.8–1.6, Q* = 1.0 (n = 110/7), sterigmata up to

ca. 110 µm long, (0.7–)1–2.3(–3.7) µm in diam.

Conidial stage common, but not always

present. Conidia difcult to distinguish from

secondary spores, and thus not measured, both of

them hyaline, smooth, thin-walled, subglobose to

ellipsoid. Conidiogenous cells hyaline, smooth,

thin-walled, globose to ellipsoid, (2.4–)3.2–

Fig. 13. Distribution of Tremella indecorata () and T.

karstenii () in Finland. The lines show the boundaries

of the biogeographical provinces.

of morphological and molecular studies and a

literature survey Chen (1998) placed all these

species in the Indecorata group. Many of their

characters are very much alike, and one might

think that the species are conspecic. There

is, however, some molecular evidence support-

ing differences. Scorzetti et al. (2002) studied

systematics of basidiomycetous yeasts based on

D1/D2 and ITS regions of ribosomal DNA, and

according to their results T. indecorata and T.

globispora are clearly distinct from each other as

well as from T. moriformis. The position of the

members of the Indecorata group is distinct on

each tree (Scorzetti et al. 2002) and differs from

the trees of Chen (1998) and Fell et al. (2000).

Further studies, including DNA techniques, are

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 421

6.5(–7.9) ¥ (1.8–)2.4–6(–6.7) µm, L = 4.7 µm, W

= 4.2 µm, Q = 0.9–2.4, Q* = 1.2 (n = 174/6).

Hyphae clamped, hyaline, smooth, (0.3–)0.9–

4(–5.9) µm in diam. (n = 104/7), thin- to slightly

thick-walled (up to 0.5 µm), oil drops common.

Haustoria originate from hyphae or conid-

iogenous cells, mostly scattered and rare, some-

times abundant or totally absent. Haustorial

mother cells hyaline, smooth, globose to ellip-

soid, (1.6–)1.8–3.5(–3.9) ¥ (1.6–)1.7–3.2(–3.4)

µm, L = 2.5 µm, W = 2.2 µm, Q = 0.9–1.4, Q*

= 1.1 (n = 39/4), each mother cell bearing one or

two haustorial laments 0.5–1 µm in diam., up

to 29 µm long, rarely branched.

Very few swollen cells observed. Vesicles

and hyphidia absent.

ecology. Tremella karstenii grows only on

Juniperus communis, and according to Torkelsen

(1978), it is always found in connection with

Colpoma juniperi. At least 16 of the 23 Finnish

collections are found on or with ascomata of C.

juniperi. The habitats of T. karstenii are mostly

coniferous forests, and it seems to prefer moist

lake- and riversides. In Finland T. karstenii is

found from April to October.

DisTribuTion anD abunDance. Collections of

T. karstenii are relatively scattered, but they come

from both southern and northern Finland (Fig.

13). We therefore presume that T. karstenii is

common and widely distributed all over the coun-

try. Tremella karstenii has a predominantly north-

ern alpine distribution. It is known from Norway,

Sweden (Torkelsen 1978, 1997), Iceland, Ger-

many, France, Austria, Switzerland, Slovakia,

Greenland, the U.S.A. (Krieglsteiner 2000), UK

(Legon et al. 2005) and Russia (Torkelsen 1978).

Records from central Europe are from the Alps or

Carpathian Mountains (Krieglsteiner 2000).

noTes. Before this study there were only

10 collections of T. karstenii from Finland. The

low number is probably due to the difculty of

nding basidiomata. They are easily overlooked

because they are small and hyaline, and shrink to

thin, almost invisible lms when dry. The easi-

est way to nd them is to search broken twigs

of Juniperus communis that lie partly buried in

mosses etc. and are thus somewhat moist.

selecTeD specimens examineD. — Finland. Varsinais-

Suomi: Merimasku, on Colpoma juniperi on Juniperus com-

munis, Karsten (S F44061). Etelä-Karjala: Hamina, Vehka-

lahti, Pyhältö, Ämmänmäki, on J. communis, 4.VII.1976

Fagerström (H). Satakunta: Merikarvia, S of Riispyy,

meadow, on C. juniperi on J. communis, 27.VIII.1969 Holm

& Holm (UPS). Etelä-Häme: Tammela, Mustiala, on C.

juniperi on J. communis, 8.IV.1869 Karsten (lectotype H).

Kainuu: Kuhmo, Ulvinsalo, Ulvinvaara, spruce-dominated

old-growth forest, on J. communis, 2005 Pippola 449 (OULU

F069356). Koillismaa: Kuusamo, Oulanka National Park,

Ampumavaara, spruce forest, on C. juniperi on J. commu-

nis, 1978 Torkelsen 106/78 (O 160931). Inarin Lappi: Inari,

Kirakkapäät, Ronkajoki, pine-dominated forest, on J. com-

munis with C. juniperi, 2005 Pippola 231 (OULU F069351).

Fig. 14. Tremella karstenii

(a–c from Karsten 1869,

d–g from Pippola 231, h

from Pippola 449). — a

and d: Spores, of which

one is forming a second-

ary spore. — b: Basidia.

— c: Hyphae. — e:

Hyphae with haustoria.

— f: Conidia or secondary

spores. — g and h: Conid-

iogenous cells.

422 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

Tremella mesenterica Retz. : Fr. (Plate 2c,

Figs. 15–16)

Kongl. Vet. Akad. Handl. 30: 249. 1769. — Tremella

mesenterica Retz. : Fr., Syst. mycol. 2 (Lundae): 214. 1822.

— Type (Wong et al. 1985): Sweden. Medelpad, Borgsjö, W

of Lillberg, on dead trunk of Salix sp., 17.VIII.1982. Strid

(neotype S).

Tremella lutescens Pers., Icon. descr. fung. 2: 33. 1798;

Syn. Meth. Fung. 2: 622. 1801. — Tremella mesenterica β

lutescens (Pers.) Pers., Mycol. Eur. 1: 100. 1822. — Tremella

lutescens Pers. : Fr., Syst. mycol. 2 (Lundae): 213. 1822.

Hormomyces aurantiacus Bonord., Handb. allgem.

Mykol.: 150. 1851, nom. anam.

Exidia candida Lloyd sensu auct.

Tremella albida Huds. : Fr. sensu auct.

Tremella candida Pers. sensu auct.

Basidiomata cream, yellow or orange-yellow,

young basidiomata cushion-like or cerebriform,

becoming folded, lobate or foliose in maturity,

some parts may be resupinate, up to 5 cm or

more in diam.

Spores hyaline to yellowish, smooth,

thin-walled, globose to subglobose, (8–)9.8–

14.9(–17.4) ¥ (7.4–)7.9–13.5(–15.6) µm, L =

12.2 µm, W = 10.5 µm, Q = 0.9–1.6, Q* = 1.2

(n = 210/7), apiculus distinctive, mostly with oil

drops, spores germinate by germ tubes or form

secondary spores.

Hymenium composed only of basidia, of

both basidia and conidia or only of conidia.

Basidia two- to four-celled, longitudinally or

occasionally obliquely septate, basally clamped,

hyaline to yellowish, smooth, mostly with oil

drops, thin-walled, globose to subglobose, some-

times wider than long, (14.1–)17.4–24.9(–25.9)

¥ (14.1–)15.6–25.4(–26.1) µm, L = 20.8 µm, W

Fig. 15. Tremella mesen-

terica (a–f from Pippola

282, g from Ulvinen 1986,

h from Korhonen 6307).

— a and g: Spores, of

which one germinates by

a germ tube and two form

secondary spores. — b:

Basidia. — c: Hyphae with

haustoria. — d: Conidia

originating from conidiog-

enous cells. — e: Various

swollen cells. — f: Vesi-

cles. — h: Hyphidia.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 423

= 20.2 µm, Q = 0.8–1.6, Q* = 1.0 (n = 110/7),

sterigmata up to 96 µm long, (1.9–)2–4.1(–4.4)

µm in diam., occasionally apically swollen up to

6.4 µm in diam., rarely branched.

Conidia hyaline to pale yellow, smooth,

thin-walled, subglobose to ellipsoid, (1.8–)2.2–

4.5(–5.9) ¥ (1.6–)1.8–3.8(–4.2) µm, L = 3.3 µm,

W = 2.6 µm, Q = 0.8–2.5, Q* = 1.3 (n = 210/7),

originating from conidiogenous cells.

Subterminal and terminal swollen cells abun-

dant close to the substrate, seldom elsewhere,

hyaline, smooth, globose, ellipsoid, citriform

or oblong, (4.5–)7.3–26.2(–41.4) ¥ (4–)5.5–

21.6(–29.4) µm, L = 16.2 µm, W = 11.3 µm, Q

= 0.9–4.4, Q* = 1.5 (n = 127/7), thin- to thick-

walled (up to 1.5 µm), occasionally short-stalked

(up to 6.4 µm in length). Swollen cells often

difcult to distinguish from vesicles and proba-

sidia.

Vesicles rare or common close to the sub-

strate, hyaline, globose, ellipsoid, ovoid, cit-

riform or oblong, (9.6–)10.4–31.6(–37.2) ¥

(6.5–)9.4–27.2(–37.6) µm, L = 20.5 µm, W =

17.1 µm, Q = 0.7–3.7, Q* = 1.3 (n = 96/7),

extremely thick walled (1.3–4.2 µm), contents

frequently oily and yellowish. Often difcult to

distinguish from swollen cells.

Hyphae clamped, hyaline to yellowish,

smooth, (0.6–)1.4–3.8(–5.6) µm in diam. (n =

119/7), mostly thin- or slightly thick-walled (up

to 0.4 µm), occasionally very thick-walled (up to

1.6 µm), oil drops common.

Hyphidia rare and scattered, usually unclear,

not measured.

Haustoria abundant close to the substrate,

originating from hyphae and sometimes from

swollen cells or vesicles. Haustorial mother cells

hyaline to yellowish, smooth, thin-walled, glo-

bose, ellipsoid or oblong, (2–)2.3–5.9(–6.4) ¥

(1.8–)2–3.9(–4.9) µm, L = 3.7 µm, W = 2.7 µm,

Q = 0.8–3.0, Q* = 1.5 (n = 106/7), each mother

cell usually bearing many, frequently branched

haustorial laments about 0.5 µm in diam., up to

30 µm long.

ecology. According to the literature (Zug-

maier et al. 1994, Roberts 1995, Torkelsen 2005)

T. mesenterica is a parasite of Peniophora spe-

cies and possibly some other closely related

genera (Zugmaier & Oberwinkler 1995, Tor-

kelsen 2005). In Finland T. mesenterica is noted

together with the following Peniophora species:

P. incarnata (4 specimens studied), P. limitata (4

specimens studied), P. violaceolivida (1 speci-

men studied) and Peniophora sp. (1 specimen

studied).

Tremella mesenterica occurs in all kind of

habitats: deciduous, coniferous and herb-rich

forests, thickets, wooded pastures, gardens and

other cultural environments. In Lapland it is

common in mountain birch forests. Basidiomata

of T. mesenterica are found on deciduous wood

(Table 2) all year round, most frequently from

August to November. It is one of the edible spe-

cies in the genus (e.g. Gorter & Eicker 1988,

Adhikari & Durrieu 1996, Boa 2004).

DisTribuTion anD abunDance. Tremella

mesenterica seems to be the most common Finn-

Fig. 16. Distribution of Tremella mesenterica () in

Finland. The lines show the boundaries of the biogeo-

graphical provinces.

424 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

ish Tremella species, followed by T. foliacea.

The bright colour may cause some misinterpreta-

tion because amateurs often bring the conspicu-

ous and attractive basidiomata to the herbaria.

Tremella mesenterica is found almost through-

out Finland (Fig. 16). Tremella mesenterica

is a cosmopolitan species, known from North

America (e.g. Coker 1920, Olive 1946b) as

well as from Central and South America (Lowy

1971), Australia (Wojewoda 1981, Roberts 1995,

Krieglsteiner 2000), New Zealand (McNabb

1966), Africa (Wojewoda 1981, Gorter & Eicker

1988, Roberts 1995, Krieglsteiner 2000), Asia

(Kobayasi 1939, Roberts 1995, Adhikari & Dur-

rieu 1996, Teng 1996, Chen 1998, Krieglsteiner

2000) and Europe (e.g. Jülich 1984, Torkelsen

1997, Krieglsteiner 2000).

noTes. The yellow, folded to foliose and

comparatively large basidiomata of T. mesente-

rica are usually easily identied. Because of the

variation in colour and the presence of conidia, T.

mesenterica is commonly considered to include

two or more separate species. However, the

variability is attributed to normal developmental

changes and responses to different light condi-

tions (Wong et al. 1985).

Tremella aurantia is another large and yellow

Tremella species known from Europe. According

to Roberts (1995), it is easily distinguished from

T. mesenterica by its substantially smaller spores

(5.5–9 ¥ 4.5–7 µm), smaller basidia (ca. 9–13

µm wide), and by the presence of host hyphae in

subhymenium and context. Moreover, the host

species separate these two taxa: T. aurantia para-

sitizes Stereum hirsutum whilst T. mesenterica

parasitizes Peniophora species (Roberts 1995).

So far, T. aurantia has not been found either in

Norway (Torkelsen 2005) or in Finland, and it

seems that the distribution of T. aurantia does

not reach the Nordic countries.

selecTeD specimens examineD. — Finland. Åland:

Lemland, Flaka, Björkö, herb-rich forest, on fallen Fraxi-

nus excelsior with Peniophora limitata, 2005 Pippola 282

(OULU F069340). Uusimaa: Espoo, Espoonkartano, on twig

of deciduous tree, 1984 Korhonen 6307 (H). Etelä-Häme:

Jaala, Kimola Canal, herb-rich forest, on cut log of Popu-

lus tremula, 1981 Haikonen 1942 (H). Keski-Pohjanmaa:

Raahe, Pikku-Kraaseli, birch forest, on Salix, 1993 Särkkä

169 (OULU F025690). Oulun Pohjanmaa: Haukipudas,

Niemeläntörmä, on Alnus, 1986 Ulvinen (OULU F068541).

Kainuu: Kuhmo, Elimyssalo, Riihipuro, peaty brookside, on

living Alnus incana, 2005 Pippola 439 (OULU F069529).

Inarin Lappi: Utsjoki, Rassijoki, on twig of Betula, 1960

Kallio (TUR 28172).

Tremella mycetophiloides Kobayasi (Plate

2d, Figs. 17–18)

Sci. Rep. Tokyo Bunrika Daigaku B 4: 13. 1939. — Type:

Japan. Prov. Kôzuke, Konsei Pass, singly or gregariously

on hymenium of living Aleurodiscus sp., on decaying trunks

of Abies veitchii Lindley, 13.VII.1938 Y. Kobayasi (type TI,

probably destroyed). — neoType (Bandoni & Ginns 1993):

Japan. Gunma Prefecture, Konsei Tooge, mainly on the west

side of the highway leading through the pass, on Aleurodis-

cus grantii on dead attached lower branches of Abies veitchii,

2.V.1986 R. J. Bandoni 7548 (DAOM).

Tremella mycophaga G.W. Martin, Mycologia 32: 686.

1940. — Type: Canada. Ontario, Algonquin Park, on Aleuro-

discus amorphus on Abies balsamea, 18 Sep. 1938 Univer-

sity of Toronto 13421 (holotype TRTC). — Synonymized by

Bandoni and Ginns (1993).

Basidiomata resupinate, cushion-like or

spherical, up to 2 mm in diam., brownish orange

when dry, translucent yellowish to pale rose

when soaked.

No spores seen, and only one mature, two-

celled, longitudinally stalked, globose basidia

observed, 10.2 ¥ 11.8 µm, single sterigma 16.1

µm long, 1.9 µm in diam. Probasidia hyaline to

yellowish, smooth, thin-walled, basally clamped,

subglobose, 10.2–14.3 ¥ 7.6–10.2 µm, L = 12.2

µm, W = 9.1 µm, Q = 1.1–1.8, Q* = 1.3 (n =

15/1).

Many basidiomata consist mainly of conid-

iogenous cells and dikaryotic and monokaryotic

conidia. Conidiogenous cells hyaline to yellow-

ish, smooth, thin-walled, globose to ellipsoid,

(6–)6.1–9.1(–9.2) ¥ (5.8–)5.9–8.4(–8.5) µm, L

= 7.7 µm, W = 7.1 µm, Q = 0.9–1.5, Q* = 1.1

(n = 26/1). Dikaryotic conidia smooth, hyaline

to yellowish, thin-walled, globose, ellipsoid or

oblong, (4.4–)4.5–6.8(–7.6) ¥ (3.3–)3.4–4.4 µm,

L = 5.8 µm, W = 3.9 µm, Q = 1.0–2.2, Q* = 1.5

(n = 30/1). Conidiogenous cells and dikaryotic

conidia form monokaryotic conidia via bud-

ding. Monokaryotic conidia smooth, hyaline,

thin-walled, subglobose to ellipsoid, (1.9–)2–

2.5(–2.6) ¥ 1.6–2.3(–2.5) µm, L = 2.2 µm, W =

1.9 µm, Q = 1.0–1.4, Q* = 1.2 (n = 30/1).

Hyphae clamped, smooth, hyaline to yellow-

ish, irregular, (1.1–)1.6–6.4(–6.6) µm in diam.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 425

(n = 24/1), thin- to slightly thick-walled (up to

0.4 µm).

Haustoria abundant. Haustorial mother cells

smooth, hyaline to yellowish, thin-walled, sub-

globose to ellipsoid 2.4–5.5 ¥ 2–3.7 µm, L =

3.8 µm, W = 2.6 µm, Q = 1.1–2.0, Q* = 1.6 (n

= 17/1), each mother cell bearing one or a few

haustorial laments up to 1.5 µm in diam. and 56

µm long, sometimes branched.

Swollen cells, vesicles and hyphidia absent.

ecology. In Finland T. mycetophiloides is

found on Aleurodiscus amorphus. In addition, it

is reported on Aleurodiscus grantii (Bandoni &

Ginns 1993) and Phanerochaete tamariciphila

(Dueñas 1996). Aleurodiscus amorphus and A.

grantii occur typically on dead branches of Abies

spp. and occasionally on species of Picea, Pinus,

Pseudotsuga and Tsuga (Ginns 1982); P. tama-

riciphila occurs on Tamarix spp. (Boidin et al.

1993). In Finland A. amorphus is found on Abies

sibirica, cultivated Abies sp., Picea abies and

Picea sp. (Kotiranta & Saarenoksa 2000). The

single Finnish collection of T. mycetophiloides is

from October.

DisTribuTion anD abunDance. Findings of the

host A. amorphus are fairly recent and few (Koti-

ranta & Saarenoksa 2000), and it is not surpris-

ing that T. mycetophiloides is also rare. Further

studies are needed to reveal the distribution and

abundance of T. mycetophiloides. At present it

Fig. 17. Tremella myce-

tophiloides (from Kotiranta

11860). — a: Probasidia

and the single mature

basidium (right). — b:

Hyphae with haustoria, of

which one is attached to

the host hyphae (arrow).

— c: Hyphae. — d: Coni-

diogenous cells, dikaryo-

tic conidia and formation

of monokaryotic conidia

(arrows).

Fig. 18. Distribution of Tremella mycetophiloides ()

and T. obscura () in Finland. The lines show the

boundaries of the biogeographical provinces.

426 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

is impossible to evaluate whether the species is

threatened or not. Worldwide T. mycetophiloides

seems to be restricted to the Northern Hemi-

sphere, as are its hosts (Ginns 1982, Boidin et al.

1993). In Europe it is reported e.g. from Norway

(Torkelsen 1968), Denmark (Christiansen 1954),

Germany (Bandoni & Ginns 1993), Spain

(Dueñas 1996), Sweden, Austria, Czech Repub-

lic, Italy, Macedonia (Pilát 1953) and Poland

(Wojewoda 1981), in Asia from Japan (Koba-

yasi 1939, Bandoni & Ginns 1993), Korea and

Taiwan (Bandoni & Ginns 1993), and in North

America both from Canada (Martin 1940, Ban-

doni & Ginns 1993) and the U.S.A. (Bandoni &

Ginns 1993).

noTes. Small, pale basidiomata found on

Aleurodiscus spp. characterize T. mycetophi-

loides. In addition, swellings in the hyphae,

which make the hyphae look irregular, and the

commonness of the conidial stage are typical.

Conidia are intermixed with basidia or alone in

the hymenium, and occurrence of the monokary-

otic conidia, rst observed by Chen (1998),

seems to be characteristic.

The same specimen was studied by Kotiranta

and Saarenoksa (2000). They managed to see one

broadly ellipsoid spore, 6 ¥ 5.5 µm, and obtained

slightly larger measures of probasidia, 15–17 ¥

(9–)11.5–13 µm. Other measurements of T. myce-

tophiloides are given for instance in the exhaus-

tive paper by Bandoni and Ginns (1993).

Tremella simplex H.S. Jacks. & G.W. Martin

is similar to T. mycetophiloides, but lacks clamps

and has consistently two-celled mature basidia

(Martin 1940).

specimen examineD. — Finland. Varsinais-Suomi:

Karjaa, Mustio manor house park, on Aleurodiscus amorphus

on dead branches of living cultivated Abies sibirica, 1994

Kotiranta 11860 (H, H.K.).

Tremella obscura (L.S. Olive) M.P. Christ.

(Figs. 18–19)

Friesia 5: 62.1954. — Tremella mycophaga G.W. Martin

var. obscura L.S. Olive, Mycologia 38: 540. 1946. — Types:

U.S.A. Georgia, Athens, deciduous woods on the University

of Georgia Campus, in the hymenium of Dacrymyces minor

on decorticated frondose wood, 25.X.1945 Lindsay S. Olive

(syntypes ISC, K(M) 36761) and in the hymenium of D.

deliquescens on cedar, 23.X.1945, Lindsay S. Olive (syntype

not located).

Basidiomata not macroscopically visible, in

the host hymenium.

Spores smooth, hyaline, thin-walled, globose

to ellipsoid, (4.3–)4.9–9(–10.4) ¥ (3.9–)3.9–

8.2(–9.8) µm, L = 7.2 µm, W = 6.3 µm, Q =

0.9–1.7, Q* = 1.2 (n = 95/4), apiculus usually

distinctive, oil drops common, spores germinate

by germ tubes. Conidia often resemble spores.

Basidia very rarely stalked, 3.5–8.4 ¥ 2.4–4.4

µm (n = 3/1), basally clamped, two-celled, lon-

gitudinally or occasionally obliquely or trans-

versely septate, smooth, hyaline to yellowish,

globose to ellipsoid, (8.4–)9.6–15.3(–16.3) ¥

(8–)9.6–14(–15.3) µm, L = 12.3 µm, W = 11.6

µm, Q = 0.8–1.7, Q* = 1.1 (n = 65/4), sterig-

mata up to 65 µm long, (1.9–)2–4.3(–6.1) µm

in diam., occasionally branched, small oil drops

common.

Conidial stage usually abundant. Conidia and

conidiogenous cells (not measured separately)

smooth, hyaline, globose, ellipsoid or oblong,

(3.3–)4.1–7.8(–8.4) ¥ (2.2–)2.5–5.7(–6) µm, L =

5.8 µm, W = 3.8 µm, Q = 1.0–2.6, Q* = 1.5 (n =

101/4), thin- to fairly thin-walled (up to 0.4 µm),

oil droplets common.

Hyphae clamped, smooth, hyaline, 1.6–

3.7(–4.3) µm in diam. (n = 58/4), thin- to rela-

tively thin-walled (up to 0.4 µm), small oil drops

common.

Haustoria abundant. Haustorial mother cells

smooth, hyaline, thin-walled, globose, subglo-

bose or ellipsoid, often wider than long, 1.8–

3.8(–4) ¥ (1.8–)1.9–3.9(–4.2) µm, L = 2.8 µm, W

= 2.7 µm, Q = 0.5–1.9, Q* = 1.1 (n = 51/4), each

mother cell bearing one haustorial lament, less

than 1 µm in diam., up to ca. 42 µm long, rarely

branched.

Swollen cells, vesicles and hyphidia absent.

ecology. Tremella obscura grows in the

hymenium of Dacrymyces species. It was origi-

nally described as parasitic on D. minor and D.

deliquescens. Finnish records are from D. lacry-

malis and D. stillatus. Tremella obscura can be

found whenever basidiomata of Dacrymyces spe-

cies are produced. Since Finnish specimens were

collected in February, May, July and August,

they can probably be found all year round.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 427

DisTribuTion anD abunDance. So far, there

are only four collections from Finland (Fig.

18). Because they are from different parts of the

country and possible hosts are common, we pre-

sume T. obscura to be a common species in the

whole Finland. Worldwide distribution is known

to extend to the U.S.A. (Olive 1946b, 1946a),

Denmark (Christiansen 1954) and UK (Roberts

2007). In addition, possible records are known

e.g. from Norway (Torkelsen 1968), Spain, Por-

tugal (Dueñas 2001) and Germany (Krieglsteiner

2000).

noTes. Tremella obscura is the rst described

species among the four known Tremella species

parasitizing Dacrymyces species. It has clamped

hyphae, mainly stalkless basidia, comparatively

short sterigmata, and thin-walled conidia which

do not arise from clamp connections. Tremella

obscura is known to have two- to four-celled

basidia (Olive 1946b, Christiansen 1954), but in

Finnish collections only two-celled basidia were

observed.

Compared with T. penetrans, T. obscura has

smaller spores and basidia and substantially

shorter sterigmata. It is not clearly distinct from

Tremella occultifuroidea Chee J. Chen & Oberw.

reported from Taiwan, but according to the origi-

nal description (Chen et al. 1999), T. occulti-

furoidea has thick-walled, dikaryotic conidia,

somewhat larger spores and remarkably long

sterigmata. T. giraffa differs from T. obscura in

its typical characters discussed earlier, but the

characters may sometimes overlap.

In addition to these other Tremella species

occurring in the hymenium of Dacrymyces spe-

cies, T. obscura is often confused with Tremella

caloceraticola Hauerslev which parasitizes

Calocera spp. as well as with the conidial stage

of Occultifur internus, an auricularioid parasite

of Dacrymyces spp. As noted by Oberwinkler

(1990), the description of T. obscura by Chris-

tiansen is based on material representing two dif-

ferent taxa: one with conidiophores of O. inter-

nus and another with tremellaceous basidia of

T. obscura, whereas the original description and

illustration by Olive (1946b) only have charac-

ters typical of T. obscura. Because Christiansen’s

description is misleading, it is not surprising that

many of the reports of T. obscura are actually

reports of the conidial stage of O. internus, or a

mixture of these two species.

specimens examineD. — Finland. Åland: Finnström,

Husö, deciduous forest, in Dacrymyces stillatus on cut top

of Alnus glutinosa with Stereum hirsutum, 2005 Pippola

292 (OULU F069349). Satakunta: Kankaanpää, Venesjärvi,

Metsäkulma, in Dacrymyces lacrymalis on Alnus, 26.II.1936

Laurila (H). Oulun Pohjanmaa: Tyrnävä, Keeterinkangas,

deciduous forest, in D. lacrymalis on twig of Salix, 1972

Ulvinen (OULU F044298). Inarin Lappi: Utsjoki, Paistun-

turit, tributary of Mavvna-avosi, mountain birch forest, in

D. lacrymalis on twig of Salix caprea, 2005 Pippola 394

(OULU F069347).

Tremella phaeophysciae Diederich & M.S.

Christ. (Plate 2e, Figs. 20–21)

in Diederich, Bibl. Lichenol. 61: 142. 1996. — Type: Den-

Fig. 19. Tremella obscura

(a–d from Ulvinen 1972,

e from Pippola 394) .

— a and e: Spores. — b:

Basidia at different stages

of development. — c:

Hyphae with haustoria, of

which one is attached to

the host hyphae (arrow).

— d: Conidia and conid-

iogenous cells.

428 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

mark. Zealand, Jungshoved, on Phaeophyscia orbicularis,

12.VIII.1966 M. S. Christiansen 2351 (holotype C; isotype

herb. Diederich).

Inducing olive brown, reddish brown, brown

or blackish brown, resupinate or somewhat

spherical or elliptical galls on the host thallus,

0.1–1.5 mm in diam., surface gelatinous, smooth

or slightly wrinkled, interior consisting of the

host hyphae.

Spores smooth, hyaline to pale brown, thin-

to fairly thin-walled (under 0.2 µm), globose,

mostly wider than long, (4.2–)5–7.8(–9.1) ¥

(4.5–)5.7–8.1(–9.4) µm, L = 6.3 µm, W = 7 µm,

Q = 0.7–1.1, Q* = 0.9 (n = 210/9), distinct apicu-

lus occasionally refractive.

Basidia two-celled, rarely basally clamped,

smooth, hyaline to pale brown, typically trans-

versely, rarely obliquely or almost longitudi-

nally septate, ellipsoid or irregularly oblong,

(11.1–)15–27.7(–31) ¥ (4.3–)5.9–9.5(–11.1) µm,

L = 21 µm, W = 7.7 µm, Q = 1.2–5.7, Q* =

2.8 (n = 134/9), sterigmata up to 27 µm long,

(1.2–)1.9–4.9(–6.2) µm in diam.

Conidial stage, especially asteroconidia,

common, but not always present. Conidia

smooth, hyaline to pale brown, thin- to fairly

thin-walled, globose to ellipsoid, (2.2–)2.5–

5.1(–5.4) ¥ (1.8–)2.1–4(–4.2) µm, L = 3.9 µm,

W = 3.1 µm, Q = 1.0–2.0, Q* = 1.3 (n =

42/4). Conidiogenous cells very unclear in some

specimens and clear and abundant in others,

smooth, hyaline, thin- to relatively thin-walled,

irregularly oblong, (11.8–)12.2–25.2(–30.2) ¥

(2.1–)2.5–5.3(–5.7) µm, L = 18.4 µm, W = 3.9

µm, Q = 2.9–9.3, Q* = 5.0 (n = 40/6), usually

with small tubes in apices (measures incl. tubes).

Four-armed asteroconidia absent in some basidi-

omata, usually abundant when present. Asteroco-

nidia smooth, hyaline to pale brown, thin-walled,

(4.1–)7.8–16(–21) µm in diam., individual arms

(0.8–)2.1–8.4(–11) µm long (n = 168/6).

Hyphae mostly simple-septate, infrequently

clamped, (1.1–)1.8–5(–7) µm in diam. (n =

139/9), thin- to thick-walled (up to 1 µm).

Haustoria few or abundant, originating from

hyphae or spores. Haustorial mother cells basally

clamped, smooth, hyaline to pale brown, thin-

walled, globose, ellipsoid or oblong, (2.1–)3.1–

5.3(–6.2) ¥ (1.8–)2–4.1(–4.3) µm, L = 4.2 µm,

W = 3 µm, Q = 0.8–2.9, Q* = 1.5 (n = 131/9),

each mother cell bearing one or two haustorial

laments under 1 µm in diam. and 18 µm long,

rarely slightly branched.

Swollen cells, vesicles and hyphidia absent.

ecology. In Finland T. phaeophysciae is

commonly found on Phaeophyscia orbicularis

(53 specimens studied) and rarely on P. endo-

phoenicea (2 specimens studied) and P. nigri-

cans (1 specimen studied). As far as we know,

this is the rst record of host species other than

P. orbicularis. Habitats of Phaeophyscia species

are mostly parks, yards, etc.

Fig. 20. Tremella phaeo-

physciae (a–f from Hausen

1927 (OULU F073618),

g–h from Hinneri 1970).

— a: Spores, of which

one bears a haustorium.

— b and g: Basidia at dif-

ferent stages of develop-

ment. — c: Hyphae with

haustoria. — d: Conidia.

— e: Conidiogenous cells.

— f: Asteroconidia. — h:

Clamped hyphae.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 429

DisTribuTion anD abunDance. Collections are

concentrated in southern Finland and the north-

ernmost collections originate from Lapua in west-

ern Finland (Fig. 21). Because P. orbicularis and

P. nigricans occur almost throughout the country

(Vitikainen et al. 1997), the distribution of T.

phaeophysciae may be much wider than currently

known. It is a common and widespread species in

Europe (Diederich 1996), but known only from a

few localities in North America (Diederich 2003).

This is the rst record from Finland.

noTes. Two-celled basidia with a transverse

septum are characteristic of T. phaeophysciae.

When the conidial stage is dominant, typically

abundant asteroconidia are found, and both the

spores and the basidia are rare. We were able to

nd only olivaceous, reddish or blackish brown

galls induced by T. phaeophysciae. According

to Diederich (1996) galls will later develop into

convex, gelatinous and usually wrinkled basidi-

omata.

It is noteworthy that KOH occasionally tinges

the microstructures purple, especially those of

the host, but to some extent also those of T.

phaeophysciae.

Tremella cladoniae is microscopically very

similar to T. phaeophysciae, but they are dis-

tinguished by the different hosts, T. cladon-

iae occurring on Cladonia spp. (see Diederich

1996). They may also differ in colour. According

to Diederich (1996) basidiomata of T. cladoniae

are more pinkish brown.

selecTeD specimens examineD. — Finland. Varsinais-

Suomi: Kisko, Kärkelän ruukki, on Phaeophyscia orbicu-

laris on Fraxinus excelsior, 1989 Laine (TUR 178741).

Uusikaupunki, N of Kasarminlahti, park, on P. orbicula-

ris on Acer platanoides, 1970 Hinneri (OULU F073616).

Satakunta: Pori, old graveyard, on P. endophoenicea on Acer,

1925 Hausen (TUR-V 08367). Pori, on P. orbicularis on

Acer, 1927 Hausen (OULU F073618). Pori, on P. orbicularis

on Acer, 1927 Hausen (OULU F073615). Etelä-Häme: Hat-

tula, Lepaa, on P. orbicularis on Sorbus aucuparia, 1935

Räsänen (TUR 178747). Etelä-Pohjanmaa: Lapua, on P.

orbicularis on S. aucuparia, 1918 Räsänen (TUR 178725).

Lapua, churchyard, on P. nigricans on Populus tremula,

1920 Räsänen (TUR-V 08471). Pohjois-Karjala: Joensuu,

Rantakatu, on P. orbicularis on S. aucuparia, 1948 Leppälä

(TUR 178735).

Tremella polyporina D.A. Reid (Figs. 22–23)

Trans. Br. Mycol. Soc. 55: 416. 1970. — Type: UK. Scot-

land, Glasgow, Cadzow Park, in the tubes of Tyromyces

lacteus which was growing either on Ulmus or on Fraxinus,

14.IX.1959, D. A. Reid (holotype K(M) 32740).

Sebacina polyporophaga Hauerslev, Friesia 11: 106.

(1976)1977. — Type: Denmark. Sealand, Tokkekøb Hegn,

on Tyromyces caesius, 28.IX.1969, K. Hauerslev 3454 (holo-

type C). — Synonymized by Wojewoda (1981).

Basidiomata not macroscopically visible, in

the host hymenium.

Spores smooth, hyaline, thin-walled, globose

to subglobose, (3.9–)4–6.2(–7.1) ¥ (3.3–)3.8–

6.1(–7) µm, L = 5.3 µm, W = 5 µm, Q = 0.8–1.3,

Q* = 1.1 (n = 120/4), apiculus distinctive, oil

drops common.

Basidia two- or four-celled, smooth, thin-

walled, longitudinally septate, basally clamped,

globose, subglobose or ellipsoid, (6.2–)7.1–

Fig. 21. Distribution of Tremella phaeophysciae () in

Finland. The lines show the boundaries of the biogeo-

graphical provinces.

430 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

13.2(–14.1) ¥ (6–)7–13.3(–13.9) µm, L =

10.4 µm, W = 10.1 µm, Q = 0.8–1.4, Q* =

1.0 (n = 63/4), sterigmata up to 17 µm long,

(0.9–)1.1–2.5(–3.5) µm in diam., usually nar-

rowing towards apices, occasionally branched,

oil drops common.

Conidial stage common. Conidia smooth,

hyaline, globose, ellipsoid or oblong, (2–)2.7–

6.2(–7.6) ¥ (1.3–)1.8–3.1(–3.7) µm, L = 4 µm,

W = 2.3 µm, Q = 1.0–3.3, Q* = 1.8 (n =

120/4), thin- to fairly thin-walled (up to 0.3 µm),

originating from conidiogenous cells and pos-

sibly infrequently from hyphal tips, oil droplets

common (seen as black spots).

Hyphae clamped, smooth, (1–)1.8–4(–5) µm

(n = 62/4), thin- to thick-walled (up to 0.8 µm),

oil drops common.

Haustoria few or abundant. Haustorial mother

cells smooth, hyaline, thin-walled, globose to

ellipsoid, 2–3.9(–4) ¥ (1.8–)1.9–3.2(–3.3) µm,

L = 2.6 µm, W = 2.3 µm, Q = 0.7–1.8, Q* = 1.1

(n = 55/4), each mother cell bearing one to three

haustorial laments 0.5–1 µm in diam., up to 20

µm long, rarely slightly branched.

Swollen cells, vesicles and hyphidia absent.

ecology. Tremella polyporina is found espe-

cially in the hymenium of Postia species, but it

may not be highly host-specic. It might rather

be a generalist occuring on various polypore

species. Finnish specimens were collected in

August, September or October.

Fig. 22. Tremella polypo-

rina (a–d from Kytömäki

3/05, e–g from Risunen

& Jakobsson 1877). — a

and e: Spores. — b and

f: Basidia. — c: Hyphae

with haustoria. — d and

g: Conidia originating from

conidiogenous cells.

Fig. 23. Distribution of Tremella polyporina () and T.

ramalinae () in Finland. The lines show the bounda-

ries of the biogeographical provinces.

ANN. BOT. FENNICI Vol. 45 • The genus Tremella in Finland 431

DisTribuTion anD abunDance. Four Finnish

collections derive from different parts of the

country (Fig. 23). Collections are so widely

distributed and the hosts are so common that we

can presume T. polyporina to be a common spe-

cies in the whole Finland. After being described

from Scotland (Reid 1970), T. polyporina has

been reported from several European countries,

e.g. Sweden (Hansen & Knudsen 1997), Norway

(Ryvarden 1998), Germany (Krieglsteiner 2000),

Czech Republic, Italy (Pouzar & Vampola 1993),

and Spain (Dueñas 1997). It is also known from

the U.S.A. (Setliff 1982) and Canada (Koske

1972). This is the rst Finnish record.

noTes. In addition to T. polyporina, Tremella

telleriae M. Dueñas occurs in the hymenium of

Postia species (Dueñas 2001). If it is distinct

from T. polyporina (Roberts 2007), it differs in

having larger basidia (Dueñas 2001) and prob-

ably also longer sterigmata and more irregular

hyphae. Spores are similar (Dueñas 2001).

A specimen found in the hymenium of Antro-

dia sinuosa has remarkably smaller spores (Table

1) and basidia than other Finnish T. polyporina

specimens, but does not differ in other respects.

Because there is so far only a single collection,

it is not considered as a new Tremella species.

Further collections combined with DNA tech-

niques are needed to understand the taxonomic

position of the specimen in A. sinuosa as well as

the taxonomy of all Tremella species occuring in

the hymenium of polypores.

specimens examineD. — Finland. Uusimaa: Helsinki,

Vanhakaupunki, Pornaistenniemi, herb-rich forest, in Postia

caesia on Alnus incana, 2005 Kiema 176 (H & H.K.). Etelä-

Häme: Lammi, Evo, Iso-Ruuhijärvi, burned, old clear-cut

area, in Postia tephroleuca on Betula with Pycnoporus cin-

nabarinus, Polyporus brumalis, Fomes fomentarius, Fomi-

topsis pinicola, Lenzites betulinus, Piptoporus betulinus,

Trametes velutina and T. hirsuta, 2003 Miettinen 7833,3 &

Pippola (H). Tampere, Soukonvuori, old spruce-dominated

forest, in Postia sp. on fallen crown of Picea, 2005 Kytömäki

3/05 (H, H.K.). Inarin Lappi: Inari, Kakslauttanen, Kiilopää,

in Antrodia sinuosa on Pinus log, 1996 Risunen & Jakobsson

1877 (K(M) 137267).

Tremella ramalinae Diederich (Plate 2f,

Figs. 23–24)

Bibl. Lichenol. 61: 152. 1996. — Type: Mexico. Baja Cali-

fornia, near km 45 on road from San Quintin to Parador

Punta Prieta, ca. 10 km N of El Rosario, 30°8´N, 115°46´W,

on Ramalina lacera, 5.I.1989 H. Sipman 24905 (holotype B;

isotype herb. Diederich).

Basidiomata pale rose to pale brown, discoid

or cushion-like, 0.2–3 mm in diam., sometimes

basally constricted, surface smooth in young

basidiomata, wrinkled in old.

Spores smooth, hyaline, thin-walled, globose

to subglobose, (4.3–)4.5–7.3(–7.7) ¥ (4–)5.1–

7.7(–7.9) µm, L = 5.8 µm, W = 6.2 µm, Q =

0.8–1.3, Q* = 1.0 (n = 60/2), distinct apiculus

often refractive.

Basidia four-celled, smooth, hyaline, thin-

walled, oblong or ovoid, (15.3–)19.2–33.3(–33.5)

¥ (8.5–)9.4–13.6(–13.8) µm, L = 24.6 µm, W

= 10.8 µm, Q = 1.4–3.4, Q* = 2.3 (n = 30/2),

uppermost septum longitudinal or oblique, two

lower septa transverse, sterigmata up to 26 µm

long, (1.9–)2–4.2(–4.9) µm in diam., sometimes

with oil drops.

Hyphae simple-septate except the basally

clamped haustoria. Hyphae smooth, hyaline,

(1–)1.2–3.8(–4) µm in diam. (n = 29/2), mostly

thin-walled, occasionally thick-walled (up to 0.8

µm).

Haustoria abundant, originating from hyphae.

Haustorial mother cells basally clamped, smooth,

hyaline, thin-walled, globose to ellipsoid,

(3.2–)3.4–4.5(–4.7) ¥ (2.3–)2.4–4.3(–4.4) µm,

L = 3.9 µm, W = 3.7 µm, Q = 0.8–1.7, Q* = 1.1

(n = 30/2), each mother cell bearing one hausto-

rial lament under 1 µm in diam. and 8 µm long,

rarely branched.

Conidial stage, swollen cells, vesicles and

hyphidia absent.

ecology. In Finland T. ramalinae is found

on the basal parts of the thallus of Ramalina

fraxinea. It is also reported on Ramalina lacera

(Diederich 1996), but R. lacera does not exist

in Finland (Vitikainen et al. 1997). Both of the

Finnish specimens of T. ramalinae were col-

lected in July.

DisTribuTion anD abunDance. The host spe-

cies R. fraxinea is mainly found in Southwest

Finland (Vitikainen et al. 1997). Thus, distribu-

tion of T. ramalinae could be wider than cur-

rently known (Fig. 23), but further studies are

needed to reveal its distribution and abundance

and evaluate its possible Red List status.

432 Pippola & Kotiranta • ANN. BOT. FENNICI Vol. 45

Tremella ramalinae seems to be a rare spe-

cies worldwide. So far, only single specimens

are known from Sweden, Mexico (Diederich

1996), Spain (Triebel 1997), Estonia (Halonen et

al. 2000), Lithuania (Motiejūnaitė 2002), Poland

(Kowalewska & Kukwa 2003), Iran (Sohrabi &

Alstrup 2007) and California (Diederich 2003).

This is the rst Finnish record.

noTes. Tremella ramalinae is easily distin-

guished from other lichenicolous Tremella spe-

cies by its four-celled basidia where the two

lower septa are transverse and the uppermost

septum longitudinal or oblique.

specimens examineD. — Finland. Åland: Hammarland,

Kattby, on Ramalina fraxinea on Fraxinus excelsior, 1964

Haakana (OULU F073614). Saltvik, Kvarnbo, on R. frax-

inea on F. excelsior, 1914 Seppälä (OULU F073617).

Conclusions

Though knowledge of Tremella species and

their distribution and abundance in Finland is

extended through this study, there is still much

to be elucidated. There may well be more than

16 species in Finland, particularly amongst the

lichenicolous taxa. Even species new to science

could yet be found. The knowledge of distribu-

tion, abundance and ecology of some Tremella

species is still too decient to evaluate if they

are threatened in Finland. Further collections are

clearly needed.

During the study we encountered many prob-

lems which need to be solved in order to under-

stand taxonomic relationships within the genus

Tremella. For instance, the taxonomy of species

associated with pyrenomycetes and parasitic in

the hymenium of polypores or Dacrymyces spe-

cies needs clarication, and a neotype for T.

foliacea must be selected. Even though all spe-

cies in the genus Tremella have characteristics in

common, it is not clear how closely lichenicolous

and other Tremella species are related.

In addition, other heterobasidioid genera war-

rant investigation in Finland.

Acknowledgements

The study is a part of the research programme of deciently

known and threatened forest species (PUTTE) funded by the

Finnish Ministry of the Environment, project 54499 led by

Minna-Maarit Kytöviita (Oulu). The curators of the listed

herbaria and C are warmly thanked for loans and/or compari-

son material. We are grateful to Robert Bandoni (Vancouver),

Paul Diederich (Luxembourg), Peter Roberts (Kew) and

Anna-Elise Torkelsen (Oslo) for giving professional advice

during the study, to Raino Lampinen (Helsinki) for making

the distribution maps, to Teuvo Ahti (Helsinki) for guid-

ance with the nomenclature and to Matti Kulju (Oulu) for

identifying part of the accompanying species. We are much

obliged to Arto Puolasmaa (Turku) who found almost all the

T. phaeophysciae specimens and identied their host lichens.

All those who helped collect new specimens deserve our

thanks, especially Pentti Alanko (Helsinki), Pekka Halonen

(Oulu), Heini Hyvärinen (Oulu), Reetta Knuuttila (Rov-

aniemi), Otto Miettinen (Helsinki), Esteri Ohenoja (Oulu),

and Teppo Rämä (Oulu).

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