28.12.2013 Views

Four new lichen-associated Trimmatostromaspecies ... - SNL

Four new lichen-associated Trimmatostromaspecies ... - SNL

Four new lichen-associated Trimmatostromaspecies ... - SNL

SHOW MORE
SHOW LESS

Create successful ePaper yourself

Turn your PDF publications into a flip-book with our unique Google optimized e-Paper software.

<strong>Four</strong> <strong>new</strong> <strong>lichen</strong>-<strong>associated</strong> Trimmatostroma species<br />

(hyphomycetes)<br />

Paul Diederich 1 , Uwe Braun 2 , Bettina Heuchert 2 & Damien Ertz 3<br />

1<br />

Musée national d’histoire naturelle, 25 rue Munster, L-2160 Luxembourg, Luxembourg (paul.diederich@education.lu)<br />

2<br />

Martin Luther University, Institute of Biology, Department of Geobotany and Botanical Garden, Herbarium,<br />

Neuwerk 21, D-06099 Halle (Saale), Germany (uwe.braun@botanik.uni-halle.de and bettina.heuchert@botanik.<br />

uni-halle.de)<br />

3<br />

Jardin Botanique National de Belgique, Domaine de Bouchout, B-1860 Meise, Belgique (damien.ertz@br.fgov.be)<br />

Diederich P., U. Braun, B. Heuchert & D. Ertz, 2010. <strong>Four</strong> <strong>new</strong> <strong>lichen</strong>-<strong>associated</strong> Trimmatostroma<br />

species (hyphomycetes). Bulletin de la Société des naturalistes luxembourgeois 111:<br />

47-55.<br />

Abstract. <strong>Four</strong> <strong>new</strong> <strong>lichen</strong>-<strong>associated</strong> species of Trimmatostroma s. lat. are described.<br />

T. dendrographae is found on Dendrographa in California (U.S.A.), T. hierrense on cf.<br />

Arthonia endlicheri in El Hierro (Canary Islands), T. lecanoricola on saxicolous Lecanora<br />

in Peru; T. quercicola develops over bark of Quercus and is facultatively <strong>lichen</strong>icolous on<br />

degenerate <strong>lichen</strong> thalli in Belgium and Luxembourg. A key to the known <strong>lichen</strong>icolous<br />

species of Trimmatostroma is provided.<br />

Key words. Anamorphic fungi, Intra<strong>lichen</strong>, <strong>lichen</strong>icolous, Taeniolella.<br />

1. Introduction<br />

The genus Trimmatostroma Corda (1837),<br />

with T. salicis Corda as type species, was<br />

introduced for bark-inhabiting saprobic<br />

hyphomycetes. This genus is characterized<br />

by having simple, little differentiated<br />

conidiophores and conidiogenous<br />

cells as well as polymorphous conidia<br />

formed in irregular basipetal chains (Ellis<br />

1971, 1976). T. <strong>lichen</strong>icola M.S. Christ. &<br />

D. Hawksw. (in Hawksworth 1979) was<br />

the first <strong>lichen</strong>icolous species assigned to<br />

this genus. Hawksworth & Cole (2002)<br />

revised and reassessed T. <strong>lichen</strong>icola and<br />

similar <strong>lichen</strong>icolous hyphomycetes and<br />

introduced the <strong>new</strong> genus Intra<strong>lichen</strong> D.<br />

Hawksw. & M.S. Cole, mainly differentiated<br />

from Trimmatostroma by its <strong>lichen</strong>icolous<br />

habit, an immersed mycelium, entirely<br />

immersed micronematous conidiophores,<br />

and pale, smooth-walled conidia with few<br />

septa. Based on molecular sequence analyses,<br />

Trimmatostroma proved to be very<br />

heterogeneous and must probably be confined<br />

to T. salicis and a few closely allied<br />

species phylogenetically pertaining to the<br />

Helotiales (Crous et al. 2007). Other species<br />

were transferred to the <strong>new</strong> genus Catenulostroma<br />

Crous & U. Braun (Crous et<br />

al. 2007). They are morphologically close<br />

to Trimmatostroma s. str., but belong to the<br />

Teratosphaeriaceae (Capnodiales). However,<br />

even Catenulostroma is in its current<br />

circumscription probably not monophyletic<br />

(Crous et al. 2009).<br />

During the course of monographic studies<br />

on <strong>lichen</strong>icolous Taeniolella S. Hughes<br />

species, several collections provisionally<br />

named “Taeniolella sp.” were encountered<br />

that turned out to be Trimmatostroma-like.<br />

Species of Taeniolella are easily distinguishable<br />

by having superficial, semimacronematous<br />

conidiophores, conidia formed<br />

in acropetal chains, and lacking multicellular<br />

aggregations of conidial cells, which<br />

are common in Trimmatostroma. As no<br />

sequence data are available for these species<br />

and as conidiomata are not stromatic<br />

(the type species T. salicis and related species<br />

have sporodochial conidiomata with<br />

a basal stroma), an inclusion in Trimmatostroma<br />

might be questionable. On the<br />

other hand, the formation of stromata<br />

Bull. Soc. Nat. luxemb. 111 (2010) 47


and conidiophores in sporodochia is often<br />

unreliable as distinctive generic character<br />

in hyphomycete genera. However, no<br />

other suitable genus is available. Therefore,<br />

we prefer to include the <strong>new</strong> <strong>lichen</strong>icolous<br />

taxa tentatively in Trimmatostroma<br />

s. lat. The final generic position of <strong>lichen</strong>icolous<br />

Trimmatostroma-like hyphomycetes<br />

depends on the phylogenetic affinity of the<br />

taxa concerned.<br />

Fig. 1. Trimmatostroma dendrographae. A–C, Infected thalli of Dendrographa. D, Conidiophore with chains of<br />

conidia, in water. E, Same, in KOH. F–J, Chains of conidia, in water. K, Conidiophore, in water. A: California,<br />

Monterey, Crocker Grove; B–C, F–K: holotype; D–E: Diederich 16767. Scale bars: B = 1 mm, C = 0.5 mm, D–K<br />

(same scale bar) = 10 μm.<br />

48 Bull. Soc. Nat. luxemb. 111 (2010)


2. Material and Methods<br />

The specimens examined are deposited in B,<br />

BR, HAL, NY, UCR and UPS, and in the private<br />

herbarium of P. Diederich. For microscopical<br />

examination, small fragments were<br />

examined in water, after light pressure on<br />

the cover glass, and in 5% KOH.<br />

3. Results<br />

Trimmatostroma dendrographae Diederich,<br />

Ertz, U. Braun & Heuchert sp. nov. Fig. 1<br />

MycoBank 518957.<br />

Trimmatostromatis quercicolae simile, sed catenis<br />

conidiorum macroscopice non distinctis,<br />

thallu hospitis plus minusve fuscescens, conidiis<br />

olivaceo-bruneis in aqua, intense olivaceis in<br />

KOH, 5.5–8.5 µm latis, pariete usque ad 1.2 µm<br />

crasso. Differt a Trimmatostromate salicis et T.<br />

betulino habitu <strong>lichen</strong>icola, stromatibus et sporodochiis<br />

nullis, coloniis dispersis supra thallos<br />

Dendrographae, conidiis olivaceo-brunneis, olivaceis<br />

in KOH, 0–pluriseptatis, 7–25 × 5.5–8.5<br />

µm, pariete rugoso-rimuloso.<br />

Type: U.S.A., California, Monterey Co., Carmel,<br />

Point Lobos State Reserve, 30 m alt., on Dendrographa<br />

sp., on Cupressus macrocarpa, 18 Jul.<br />

2008, P. Diederich 16789 & D. Ertz (BR – holotype;<br />

HAL, NY, UCR, herb. Diederich – isotypes).<br />

Colonies <strong>lichen</strong>icolous over the thallus of<br />

Dendrographa, usually covering the entire<br />

surface of the host thallus, blackish, macroscopically<br />

appearing as a finely granular<br />

mixture of blackish conidia and white<br />

host hyphae, the whole appearing pale to<br />

dark greyish at a low magnification (e.g.,<br />

in the field), individual chains of conidia<br />

hardly visible. Mycelium internal, hyphae<br />

sparingly branched, 2.5–3.5 µm wide, subhyaline<br />

to pale brown, septate, thin-walled,<br />

rugose. Stroma lacking. Conidiophores usually<br />

immersed, occasionally somewhat<br />

erumpent, micronematous, c. 20–70 × 2.5–4<br />

µm, hyphal filaments gradually developing<br />

into fertile threads by becoming somewhat<br />

wider and darker, with slightly thicker<br />

wall, but differentiation between hyphae<br />

and conidiophores very difficult, rugose.<br />

Conidiogenous cells integrated, terminal,<br />

monoblastic (i.e., the terminal cell functioning<br />

as conidiogenous cell), c. 5.5–9 × 3.5–4<br />

µm, conidiogenous loci undifferentiated,<br />

subtruncate. Conidia in simple or rarely<br />

branched, irregular, rarely disarticulating,<br />

basipetal chains, shape and size very variable,<br />

mostly ellipsoid-ovoid to subcylindrical<br />

or oblong, transversely 0–pluriseptate,<br />

rarely dictyosporous, ends more or less<br />

rounded, c. 7–25 × 5.5–8.5 µm, sometimes<br />

forming irregular aggregations, medium to<br />

dark brown or olivaceous-brown, distinctly<br />

olivaceous in KOH, wall 0.8–1.2 μm thick,<br />

wall rugose-rimulose.<br />

This species is distinguished from T. lecanoricola<br />

by olivaceous conidia in KOH, from<br />

T. hierrense by rugose-rimulose conidia and<br />

from T. quercicola by narrower conidia that<br />

are not visible macroscopically. Attempts to<br />

culture freshly collected material failed.<br />

Ecology, hosts and distribution: All known<br />

specimens are <strong>lichen</strong>icolous on epiphytic<br />

and saxicolous thalli of Dendrographa,<br />

including crustose, sorediate thalli that are<br />

considered to be the primary morph of the<br />

host, before the fruticose structures appear.<br />

In many specimens, fruticose thalli start<br />

growing, but are so heavily parasitized by<br />

this hyphomycete that they just reach up to<br />

1 cm in length and then die. This species is<br />

known only from the coast of California, and<br />

it appears to be rather common in localities<br />

where the host Dendrographa is present.<br />

Additional specimen examined (all on Dendrographa):<br />

USA, California, Monterey Co.:<br />

same locality as the type, on Cupressus macrocarpa,<br />

2008, Ertz 12482, 12487 & Diederich (BR);<br />

ibid., sheltered rock face near the sea, 2008, Ertz<br />

12441, 12453 & Diederich (BR); SW of Monterey,<br />

17 Mile Drive, Crocker Grove, alt. 15 m, on C.<br />

macrocarpa, 2008, Diederich 16767 & Ertz 12462<br />

(BR, herb. Diederich); along coast S of Asilomar,<br />

China Rock, alt. 5 m, on rock outcrop on the<br />

beach, 2008, Ertz 12480 & Diederich (BR).<br />

Trimmatostroma hierrense Diederich &<br />

Ertz sp. nov. Fig. 2<br />

MycoBank 518958.<br />

Differt ab omnibus speciebus Trimmatostromatis<br />

conidiis transverse septatis et parietibus conidiorum<br />

unilateraliter incrassatis et fuscatis.<br />

Type: Spain, Canary Islands, El Hierro, E of<br />

Frontera, trail between Frontera and ‘mirador de<br />

Jinama’ (part below ‘Fuente de Tincos’), 27°45’29”<br />

N, 17°59’29” W, ‘fayal brezal’ forest and rocky<br />

outcrops, on a root under a rock along trail, on cf.<br />

Bull. Soc. Nat. luxemb. 111 (2010) 49


Fig. 2. Trimmatostroma hierrense (holotype). A–B, Infected thalli of cf. Arthonia endlicheri. C, Conidia and conidial<br />

chains, in water. D–F, Same, in KOH (conidiophores visible in D). Scale bars: A = 1 mm, B = 0.2 mm, C–F = 10 μm.<br />

Arthonia endlicheri, 31 Aug. 2009, D. Ertz 13946<br />

(BR – holotype; herb. Diederich – isotypes).<br />

Colonies <strong>lichen</strong>icolous over the thallus of cf.<br />

Arthonia endlicheri, covering large parts of<br />

the host thallus, blackish, macroscopically<br />

appearing as a finely granular mixture of<br />

blackish conidia and white host hyphae, the<br />

whole appearing dark greyish to blackish at<br />

a low magnification (e.g., in the field), individual<br />

chains of conidia hardly visible. Mycelium<br />

internal, hyphae sparingly branched,<br />

2–2.5 µm wide, subhyaline to pale brown,<br />

septate, thin-walled, smooth. Stroma lacking.<br />

Conidiophores usually immersed, occasionally<br />

somewhat erumpent, micronematous, up<br />

to 40 μm long and 2.5–3.5 µm wide, hyphal<br />

filaments gradually developing into fertile<br />

threads by becoming somewhat wider and<br />

darker, but differentiation between hyphae<br />

and conidiophores very difficult, smooth.<br />

Conidiogenous cells integrated, terminal,<br />

monoblastic (i.e., the terminal cell functioning<br />

as conidiogenous cell), c. 5.5–10 ×<br />

3.5–4.5 µm, conidiogenous loci undifferentiated,<br />

subtruncate. Conidia in simple or rarely<br />

branched, irregular, often disarticulating,<br />

basipetal chains, shape and size very variable,<br />

mostly ellipsoid-ovoid to subcylindrical<br />

or oblong, often with irregular convex and<br />

concave regions, transversely 0–pauciseptate,<br />

ends more or less rounded, c. 7.5–15 × 3.5–<br />

6.5 µm, pale to medium brown, partly with<br />

thickened walls and then dark brown, olivaceous<br />

brown in KOH; wall c. 0.5 μm thick in<br />

pale areas, c. 1 μm in darker areas, smooth.<br />

Macroscopically, the <strong>new</strong> species looks very<br />

similar to Trimmatostroma dendrographae,<br />

entirely darkening large areas of the host<br />

50 Bull. Soc. Nat. luxemb. 111 (2010)


thallus. Microscopically, it is distinguished<br />

by conidia frequently with a dark and thick<br />

outer wall limited to one side of the conidium<br />

(resembling conidia of Milospium<br />

graphideorum). Furthermore, conidia are<br />

usually smoother than in T. dendrographae,<br />

best seen in KOH. Because of the unevenly<br />

pigmented conidial wall, T. hierrense might<br />

be mistaken for a species of Milospium.<br />

However, conidia of the type species, M.<br />

graphideorum (Nyl.) D. Hawksw., have a<br />

very different shape, being irregularly lobed<br />

and aseptate, and they are never catenate.<br />

Conidia of M. deslooveri Diederich & Sérus.<br />

(Sérusiaux et al. 1999) and M. planorbis<br />

(Aptroot & Sipman 2001; holotype from<br />

B examined) are strongly curved, multicellular,<br />

often with incomplete septa, and<br />

never catenate; furthermore, both species<br />

are <strong>lichen</strong>ized, not <strong>lichen</strong>icolous. The <strong>new</strong><br />

species could also be confused with Trimmatostroma<br />

scutellare (Berk. & Broome) M.<br />

B. Ellis, another species with an unevenly<br />

thickened and pigmented conidial wall,<br />

but in that species conidia are multicellular<br />

with transverse and longitudinal septa (Ellis<br />

1976).<br />

Ecology, hosts and distribution: The <strong>new</strong><br />

species covers large parts of the thallus of<br />

a white crustose <strong>lichen</strong> with Trentepohlia<br />

reacting C+ red, that probably belongs to<br />

Arthonia endlicheri, growing on the root of<br />

a tree along a trail. Infected thalli become<br />

entirely dark greyish black. This species is<br />

known only from the type locality on the isle<br />

of El Hierro (Spain, Canary Islands).<br />

Trimmatostroma lecanoricola Diederich,<br />

Etayo, U. Braun & Heuchert sp. nov. Fig. 3<br />

MycoBank 518959.<br />

Trimmatostromatis dendrographae et T. quercicolae<br />

simile, sed catenis conidiorum disarticulantium<br />

aurantiaco-brunneis in KOH, nec umquam<br />

Fig. 3. Trimmatostroma lecanoricola (holotype). A, Thallus of saxicolous Lecanora with young infection. B, Thallus<br />

with older infection. C, Chains of conidia, in water. D, Same, in KOH. E, Disarticulating chain of conidia, in water.<br />

F, Conidium in surface view, showing verrucose, mosaic-like outer layer of conidial wall, in water. Scale bars: A–B<br />

= 0.5 mm, C–F = 10 μm.<br />

Bull. Soc. Nat. luxemb. 111 (2010) 51


olivaceo-brunneis, et partibus atrioribus paginae<br />

conidiorum minoribus in conidiis vetustioribus<br />

rimulosis.<br />

Type: Peru, Dep. La Libertad, Prov. Trujillo, Cerro<br />

Malabrigo, SW of Puerto Chicama, on saxicolous<br />

Lecanora, 100–150 m alt., 9 Apr. 1981, R. Santesson<br />

& A. Tehler P128:1 (UPS – holotype).<br />

Colonies <strong>lichen</strong>icolous, over the thallus and<br />

apothecia of a saxicolous Lecanora; when<br />

young, appearing as irregular, branched,<br />

blackish filaments (representing conidial<br />

chains), soon aggregating in very dense<br />

tufts of conidial chains, the infected parts of<br />

the host becoming entirely blackish. Mycelium<br />

internal, hyphae sparingly branched,<br />

1.5–3 µm wide, subhyaline to pale brown,<br />

septate, thin-walled, smooth. Stroma lacking.<br />

Conidiophores usually immersed, occasionally<br />

somewhat erumpent, micronematous,<br />

c. 20–40 × 2–4 µm, hyphal filaments<br />

gradually developing into fertile threads<br />

by becoming somewhat wider and darker,<br />

with slightly thicker wall, but differentiation<br />

between hyphae and conidiophores very<br />

difficult. Conidiogenous cells integrated, terminal,<br />

monoblastic (i.e., the terminal cell<br />

functioning as conidiogenous cell), c. 4–7<br />

× 3–4.5 µm, conidiogenous loci undifferentiated,<br />

subtruncate. Conidia in simple or<br />

rarely branched, irregular, easily disarticulating,<br />

basipetal chains, often subspherical<br />

and 1-septate, 6.5–10 × 6–8 μm, or oblong,<br />

transversely pluriseptate to dictyosporous,<br />

c. 11–24 × 5–9 µm, medium to dark brown,<br />

orange brown in KOH, wall 0.5–1.2 µm<br />

thick, rugose-rimulose to irregularly verrucose,<br />

ends more or less rounded.<br />

This species is readily distinguished from<br />

the other taxa treated here by conidial chains<br />

becoming orange brown in KOH, never with<br />

an olivaceous tinge, and by chains of disarticulating<br />

conidia. Furthermore, when the<br />

outer layer of the conidial wall splits with<br />

age, the resulting darker patches are often<br />

smaller than in the other species, giving the<br />

conidial surface a mosaic-like appearance.<br />

Ecology, host and distribution: This species<br />

is known only from the type collection<br />

made in Peru on an unidentified saxicolous<br />

Lecanora. Infected host thalli and apothecia<br />

are damaged, suggesting that the fungus is<br />

pathogenic.<br />

Trimmatostroma quercicola Diederich, U.<br />

Braun & Heuchert sp. nov. Figs 4–5<br />

MycoBank 518960.<br />

Trimmatostromatis dendrographae simile, sed<br />

catenis conidiorum macroscopice expedite conspicuis,<br />

irregulariter reticuloideis, conidiis badiis<br />

in aqua et olivaceo-brunneis in KOH, 5–15 µm<br />

latis, pariete usque ad 2 μm crasso. Differt a T.<br />

lecanoricola catenis conidiorum raro disarticulatis,<br />

conidiis olivaceo-brunneis in KOH, et strato<br />

exteriore conidiorum obscuriore in conidiis<br />

vetustioribus rimulosis.<br />

Type: Luxembourg, Vogelsmühle, valley of Halerbaach,<br />

left side of river, on bark of Quercus, possibly<br />

<strong>associated</strong> with sterile crustose <strong>lichen</strong>s, 9 Dec.<br />

2007, P. Diederich 16723 (BR – holotype; HAL,<br />

herb. Diederich – isotypes).<br />

Colonies over bark of Quercus, frequently<br />

overgrowing and probably parasitizing<br />

degenerate crustose <strong>lichen</strong> thalli or corticolous<br />

algae, effuse, loose to dense, fumoseblack,<br />

macroscopically appearing as relatively<br />

short, prostrate, irregularly formed<br />

and branched, often agglomerated conidial<br />

chains. Mycelium internal, hyphae sparingly<br />

branched, 1–3.5 µm wide, subhyaline to pale<br />

brown, septate, thin-walled, smooth. Stroma<br />

lacking. Conidiophores usually immersed,<br />

occasionally somewhat erumpent, micronematous,<br />

c. 5–40 × 2–6 µm, hyphal filaments<br />

gradually developing into fertile threads<br />

by becoming somewhat wider and darker,<br />

with slightly thicker wall, but differentiation<br />

between hyphae and conidiophores very<br />

difficult. Conidiogenous cells integrated,<br />

terminal, monoblastic (i.e., the terminal<br />

cell functioning as conidiogenous cell), c.<br />

5–10 × 3–4 µm, conidiogenous loci undifferentiated,<br />

subtruncate. Conidia in simple<br />

or rarely branched, irregular, occasionally<br />

disarticulating, basipetal chains, shape and<br />

size very variable, globose or subglobose,<br />

0–1(–2)-septate, 4–9 µm diam., or ellipsoidovoid,<br />

subcylindrical, oblong, 1–4(–5)-septate,<br />

transversely septate to dictyosporous, c.<br />

6–30 × 5–15 µm, sometimes forming irregular<br />

aggregations, medium to dark brown,<br />

olivaceous-brown in KOH, wall thick, up to<br />

2 µm, occasionally distinctly two-layered,<br />

i.e., with a distinct, paler inner layer (but not<br />

distoseptate), wall rugose-rimulose to irregularly<br />

verrucose, ends more or less rounded.<br />

52 Bull. Soc. Nat. luxemb. 111 (2010)


Amongst the <strong>new</strong> species described here, T.<br />

quercicola is macroscopically rather distinct,<br />

as conidial chains are easily visible at a strong<br />

magnification (× 40), appearing as an irregular,<br />

branched net of relatively thick hyphae<br />

overgrowing the substratum. Microscopically,<br />

the species has much thicker conidial<br />

chains than the three others. It is chemically<br />

distinct from T. lecanoricola by the K+ olivaceous<br />

conidial chains and differs slightly from<br />

T. dendrographae in having brown conidia (in<br />

water) without a distinct olivaceous tinge.<br />

Trimmatostroma quercinum (Hoffm.) Höhn.<br />

(syn. Phragmotrichum quercinum Hoffm.)<br />

was described from rotten manufactured<br />

wood of Quercus in Germany. The species<br />

forms black, superficial, dispersed, orbicular,<br />

pulvinate stromata, and oblong, muricate,<br />

transversally septate to irregularly<br />

muriform, indistinctly catenate conidia<br />

25–30 × 12–15 μm (Migula 1921, Sutton &<br />

Pirozynski 1965). Although the type of this<br />

species has never been restudied and the<br />

species never been reported after its original<br />

description, it seems to differ clearly from<br />

the <strong>new</strong> T. quercicola by the pulvinate, stromatic<br />

conidiomata, the different ecology<br />

(rotten wood, versus bark of Quercus) and<br />

the mostly larger conidia.<br />

Ecology, hosts and distribution: The <strong>new</strong><br />

species is known from two collections on<br />

the bark of old Quercus. As conidiophores<br />

at least occasionally overgrow and possibly<br />

parasitize reduced <strong>lichen</strong> thalli, the species<br />

may be considered as facultatively <strong>lichen</strong>icolous.<br />

It is probably widespread, but overlooked<br />

in similar habitats, and currently<br />

known from Belgium and Luxembourg.<br />

Additional specimen examined: Belgium, SE of<br />

Buzenol, on bark of Quercus, possibly <strong>associated</strong><br />

with sterile crustose <strong>lichen</strong>s, 19 June 1984, P. Diederich<br />

5576 (herb. Diederich).<br />

Fig. 4. Trimmatostroma quercicola (holotype). A, Conidiophores with and without conidia. B, Plagiotropous conidial<br />

chain. C, Conidia. Scale bar = 10 µm. U. Braun del.<br />

Bull. Soc. Nat. luxemb. 111 (2010) 53


Fig. 5. Trimmatostroma quercicola. A, Overgrowing degenerate <strong>lichen</strong> thalli on Quercus bark. B, Saprobic over Quercus<br />

bark. C, Squash preparation of infected <strong>lichen</strong> thallus, in water. D, Chains of conidia, in water. E, Same, in KOH.<br />

A, C: holotype; B, D–E: Diederich 5576. Scale bars: A–B = 200 μm, C–E = 10 μm.<br />

4. Discussion<br />

Hawksworth (1979) described the <strong>lichen</strong>icolous<br />

Trimmatostroma <strong>lichen</strong>icola M.S.Christ.<br />

& D. Hawksw., intrahymenial in Candelariella<br />

vitellina. Later, Hawksworth & Cole (2002)<br />

introduced the <strong>new</strong> generic name Intra<strong>lichen</strong><br />

for this and several similar species, and<br />

no <strong>lichen</strong>icolous species was subsequently<br />

accepted in Trimmatostroma.<br />

The four <strong>new</strong> species share a large number<br />

of characters. They all have dark, micronematous<br />

conidiophores not arranged in<br />

sporodochia, lack stromatic hyphal aggregations,<br />

have integrated conidiogenous cells,<br />

and conidia produced in simple or rarely<br />

branched basipetal chains, 1–pluriseptate,<br />

irregular in size and form, with an often<br />

rough surface, the outer layer of the wall<br />

often splitting with age, resulting in pale<br />

to medium brown conidia covered by dark<br />

brown remnants of the outer layer, or unevenly<br />

pigmented. Furthermore, they all obligately<br />

or facultatively parasitize <strong>lichen</strong> thalli.<br />

Conidia are superficially similar to those of<br />

Spilodochium Syd. (Ellis 1971, 1976). However,<br />

the genus Spilodochium is easily distinguishable<br />

by its erumpent pseudoparenchymatic<br />

stromata, conidiophores arranged<br />

in sporodochia and conidia produced in<br />

branched acropetal chains. The entire habit<br />

of the <strong>new</strong> species is rather Intra<strong>lichen</strong>-like.<br />

However, species of Intra<strong>lichen</strong> have paler,<br />

smaller, smooth-walled, mostly 0–2-celled<br />

conidia, and conidiophores are usually<br />

entirely immersed in <strong>lichen</strong> thalli or apothecia,<br />

most commonly being intrahymenial,<br />

with only the mature conidia arising at the<br />

surface (Hawksworth & Cole 2002). On the<br />

other hand, the inclusion of these species<br />

in Taeniolella is no alternative, since species<br />

of this genus do not form dictyosporous<br />

conidia, and its conidial chains are acropetal.<br />

Microscopically, the <strong>new</strong> species are similar<br />

to Trimmatostroma salicis, the type species,<br />

and T. betulinum (Corda) S.Hughes, from<br />

which they are distinguished by the absence<br />

of a basal stroma and lacking pulvinate spo-<br />

54 Bull. Soc. Nat. luxemb. 111 (2010)


odochia (Crous et al. 2007). The latter two<br />

species have been shown to belong to the<br />

Helotiales (Crous et al. 2007). Many Trimmatostroma-like<br />

fungi have recently been<br />

transferred to the <strong>new</strong> genus Catenulostroma<br />

in the Teratosphaeriaceae [Capnodiales]<br />

(Crous et al. 2007), which is, however,<br />

no alternative either, since this genus is also<br />

characterized by having stromatic conidiomata.<br />

The introduction of a <strong>new</strong> genus for<br />

the <strong>new</strong> <strong>lichen</strong>icolous species might be the<br />

best solution, but in the absence of molecular<br />

data, i.e., without knowledge of the phylogenetic<br />

affinity, we prefer to assign the <strong>new</strong><br />

species tentatively to Trimmatostroma s. lat.<br />

5. Identification key for the known<br />

<strong>lichen</strong>icolous species of Trimmatostroma<br />

1. Conidia orange brown in KOH; conidial<br />

chains frequently disarticulating; outer layer of<br />

conidial wall splitting with age and then sometimes<br />

divided in many relatively small patches,<br />

looking mosaic-like (Fig. 3F); <strong>lichen</strong>icolous on<br />

saxicolous Lecanora................... T. lecanoricola<br />

1. Conidia olivaceous-brown in KOH;<br />

conidial chains rarely disarticulating; outer<br />

layer of conidial wall either smooth (Fig. 2)<br />

or splitting with age and then divided into<br />

several larger patches (Figs 1 & 5D) .......... 2<br />

2. Conidial chains easily visible macroscopically<br />

(× 40), appearing as an irregular net of<br />

blackish hyphae; conidia reddish brown in<br />

water, olivaceous-brown in KOH, 5–15 µm<br />

wide, wall up to 2 μm thick; saprobic on bark<br />

of Quercus, facultatively <strong>lichen</strong>icolous over<br />

degenerated <strong>lichen</strong> thalli ........... T. quercicola<br />

2. Conidial chains indistinct macroscopically,<br />

almost entirely blackening large parts<br />

of the host thallus; conidia olivaceousbrown<br />

in water, more intensively olivaceous<br />

in KOH, 5.5–8.5 µm wide, wall up to 1.2 μm<br />

thick; obligately <strong>lichen</strong>icolous on <strong>lichen</strong>s<br />

with Trentepohlia .......................................... 3<br />

3. Conidia frequently with a dark outer<br />

wall limited to one side of the conidium<br />

(resembling conidia of Milospium), wall<br />

smooth; <strong>lichen</strong>icolous on cf. Arthonia endlicheri<br />

.......................................... T. hierrense<br />

3. Conidia with a dark outer wall splitting<br />

with age, but more evenly surrounding the<br />

entire conidia, wall rugose; <strong>lichen</strong>icolous on<br />

Dendrographa .................. T. dendrographae<br />

Acknowledgements<br />

We are much obliged to the curators of B and<br />

UPS for the possibility to examine collections<br />

from their herbaria, and to Prof. Walter Gams for<br />

critically commenting on the manuscript.<br />

References<br />

Aptroot, A. & H. J. M. Sipman, 2001. New Hong<br />

Kong <strong>lichen</strong>s, ascomycetes and <strong>lichen</strong>icolous<br />

fungi. Journal of the Hattori Botanical Laboratory<br />

91: 317–343.<br />

Corda, A. K. J., 1837. Icones fungorum hucusque<br />

cognitorum. Vol. 1. J. G. Calve, Praha.<br />

Crous, P. W., U. Braun, J. Z. Groe<strong>new</strong>ald, 2007.<br />

Mycosphaerella is polyphyletic. Studies in<br />

Mycology 58: 1–32.<br />

Crous, P. W., C. L. Schoch, K. D. Hyde, A. R.<br />

Wood, C. Gueidan, G. S. de Hoog & J. Z.<br />

Groe<strong>new</strong>ald, 2009. Phylogenetic lineages<br />

in the Capnodiales. Studies in Mycology 64:<br />

17–47.<br />

Ellis, M. B., 1971. Dematiaceous hyphomycetes.<br />

Commonwealth Mycological Institute, Kew.<br />

Ellis, M. B., 1976. More dematiaceous hyphomycetes.<br />

Commonwealth Mycological Institute,<br />

Kew.<br />

Hawksworth, D. F., 1979. The <strong>lichen</strong>icolous<br />

hyphomycetes. Bulletin of the British Museum<br />

(Natural History), Botany 6: 183–300.<br />

Hawksworth, D. F. & M. S. Cole, 2002. Intra<strong>lichen</strong>,<br />

a <strong>new</strong> genus for <strong>lichen</strong>icolous ‘Bispora’<br />

and ‘Trimmatostroma’ species. Fungal Diversity<br />

11: 87–97.<br />

Migula, W., 1921. Fungi imperfecti: Sphaeropsidales,<br />

Melanconiales. Kryptogamen-Flora<br />

von Deutschland, Deutsch-Österreich und<br />

der Schweiz, Band III, 4. Teil, 1. Abteilung.<br />

Berlin, Hugo Bermühler Verlag.<br />

Sérusiaux, E., P. Diederich, M. Brand & P. van den<br />

Boom, 1999. New or interesting <strong>lichen</strong>s and<br />

<strong>lichen</strong>icolous fungi from Belgium and Luxembourg.<br />

VIII. Lejeunia n. S. 162: 1–95.<br />

Sutton, B.C. & K. A. Pirozynski, 1965. Notes on<br />

microfungi. II. Transactions of the British<br />

Mycological Society 48: 349–366.<br />

Bull. Soc. Nat. luxemb. 111 (2010) 55

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!