Lichenicolous Lichens and Fungi from Monfragüe National Park (Western Spain)

Abstract

Etayo, J. & Pérez-Ortega, S. 2016. Lichenicolous lichens and fungi from Monfragüe National Park (western Spain). — Herzogia 29: 315–328.
We present the first survey of lichenicolous lichens and fungi in Monfragüe National Park, Cáceres (Spain). Seven localities were visited, including the most characteristic ecosystems. Forty taxa were recorded, mostly on epiphytic lichens. Seven species are reported from the Iberian Peninsula for the first time: Acremonium subeffusum, Cornutispora intermedia, Endococcus exerrans, Pronectria subimperspicua, Roselliniella cladoniae, Tremella cladoniae and Zwackhiomyces echinulatus. The species Arthonia pepei, growing on Parmelina cryptotiliacea, is described as new to science.

Full text

Herzogia 29 (2) Teil 1, 2016: 315 –328 315
Lichenicolous lichens and fungi from Monfragüe National Park
(western Spain)
Javier Etayo* & Sergio Pérez-Ortega
Abstract: Etayo, J. & Pérez-Ortega, S. 2016. Lichenicolous lichens and fungi from Monfragüe National Park
(western Spain). – Herzogia 29: 315 –328.
We present the first survey of lichenicolous lichens and fungi in Monfragüe National Park, Cáceres (Spain). Seven
localities were visited, including the most characteristic ecosystems. Forty taxa were recorded, mostly on epiphytic li-
chens. Seven species are reported from the Iberian Peninsula for the first time: Acremonium subeffusum, Cornutispora
intermedia, Endococcus exerrans, Pronectria subimperspicua, Roselliniella cladoniae, Tremella cladoniae and
Zwackhiomyces echinulatus. The species Arthonia pepei, growing on Parmelina cryptotiliacea, is described as new
to science.
Zusammenfassung: Etayo, J. & Pérez-Ortega, S. 2016. Lichenicole Flechten und Pilze aus dem Nationalpark
Monfragüe (westliches Spanien). – Herzogia 29: 315 –328.
Die Ergebnisse einer ersten Erhebung lichenicoler Flechten und Pilze im Nationalpark Monfragüe, Cáceres (Spanien)
werden vorgelegt. Sieben Lokalitäten, welche die charakteristischsten Ökosysteme des Gebietes repräsentieren, wur-
den besucht und insgesamt 40 Taxa (die meisten davon auf epiphytischen Flechten) dokumentiert. Sieben der Arten
werden für die Iberische Halbinsel zum ersten Mal nachgewiesen: Acremonium subeffusum, Cornutispora intermedia,
Endococcus exerrans, Pronectria subimperspicua, Roselliniella cladoniae, Tremella cladoniae und Zwackhiomyces
echinulatus. Die auf Parmelina cryptotiliacea wachsende Art Arthonia pepei wird hier neu beschrieben.
Key words: Arthoniaceae, Ascomycota, Basidiomycota, Mediterranean, hyphomycetes.
Introduction
The Iberian Peninsula, and especially Spain, is among the best-studied regions in terms of
lichenicolous fungi (e.g. Calatayud & Etayo 2001, Llimona & Hladun 2001, Martínez
2002, Hafellner & Casares-Porcel 2003, Llop & Gómez-Bolea 2006, Navarro-Rosinés
et al. 2008, Pérez-Ortega & Etayo 2008, Atienza et al. 2009, Etayo 2010a, b, Zamora
et al. 2011, van den Boom & Etayo 2014). However, there are still large areas for which the
diversity of these organisms is still poorly known (Pérez-Ortega & Álvarez-Lafuente
2006), especially in southern and western regions. Monfragüe National Park is located in
western Spain, in Cáceres province. The area was declared as Natural Park in 1979 due to
the valuable and well preserved Mediterranean forest and scrub landscape. Subsequently it
was declared a special protected area (SPA) in 1994, a Biosphere Reserve in 2003, and fi-
nally a National Park in 2007. Monfragüe spans an area of 18,396 ha (between 39º52'56'' and
39º42'22''N and 6º06'26'' and 5º45'14''W) and its average elevation is 360 m a.s.l. Climate
* Corresponding author

316 Herzogia 29 (2) Teil 1, 2016
is sub-humid mediterranean and the annual rainfall is c. 791 mm (Rivas-Martínez 1987).
Forests are dominated by holm oak (Quercus rotundifolia), cork oak (Quercus suber) and
Portuguese oak (Q. faginea), with the usual presence of maples (Acer monspessulanum), ter-
ebinth (Pistacia terebinthus), and hornbeam (Celtis australis). Shrublands are also very impor-
tant with gum rockroses (Cistus ladanifer), strawberry-tree (Arbutus unedo), butcher’s broom
(Ruscus aculeatus), myrtle (Myrtus communis) or heather (Erica arborea) as the most char-
acteristic species. Calatayud et al. (2011a, b) provided the first catalogue of lichen-forming
fungi occurring at the National Park, reporting a total of 95 species, mostly macrolichens and
highly conspicuous microlichens.
During April 2015, the first author visited several localities in Monfragüe National Park in the
first attempt to catalogue the biota of lichenicolous lichens and fungi.
Material and methods
Specimens were macroscopically studied using a Meiji zoom stereomicroscope and micro-
scopically with a Nikon Eclipse 80i microscope fitted with differential interference contrast
(‘Nomarsky’). All microscopic measurements were made on hand-cut sections mounted in
water. Hand-cut sections mounted in 10 % KOH, Lugol’s iodine (I), 50 % nitric acid (N) and
phloxine were used for identification. Specimens are deposited in the private herbarium of the
first author, and some duplicates in VIT (The Natural History Museum of Alava). The holotype
of the new taxon is deposited in MAF-LICH.
Studied localities
1. Cáceres, Quercus rotundifolia dehesa in the road from Plasencia to Villarreal de San Carlos, c. 420 m a.s.l.,
13.4.2015, 39º56'42''N, 06º05'36''W.
2. Cáceres, P.N. de Monfragüe, Villarreal de San Carlos, Quercus rotundifolia dehesa with Cistus ladanifer close to
information point, 340 m a.s.l., 13.4.2015, 39º50'29''N, 06º01'51''W.
3. Cáceres, P.N. de Monfragüe, Quercus rotundifolia forest with Olea europea and Fraxinus angustifolia close to
Fuente del Francés, c. 300 m a.s.l., 13.4.2015, 39º49'46''N, 06º02'08''W.
4. Cáceres, P.N. de Monfragüe, stands with Quercus rotundifolia and Celtis australis close to Monfragüe castle and
hermitage, c. 240 m a.s.l., 13.4.2015, 39º49'42''N, 6º03'07''W.
5. Cáceres, P.N. de Monfragüe, Quercus rotundifolia dehesa with quartzitic outcrops close to Saltos del Torrejón
(Tres Caños), 230 m a.s.l., 13.4.2015, 39º50'39''N, 06º00'05''W.
6. Cáceres, P.N. de Monfragüe, Cansinas Creek, dispersed trees in the riverside near the dam, 230 m a.s.l., 13.4.2015,
39º50'35''N, 05º57'09''W.
7. Cáceres, P.N. de Monfragüe, quartzitic outcrops in Quercus rotundifolia forest close to Mirador de la Tajadilla,
240 m a.s.l., 13.4.2015, 39º50'19''N, 5º59'45''W.
The species
Numbers in italics indicate localities described above. Asterisks before taxon name indicate
lichenicolous lichenized fungi.
Abrothallus microspermus Tul.
Only found in its anamorphic state [= Vouauxiomyces truncatus (de Lesd.) Dyko & D.Hawksw.].
Previously known from Catalonia (Glenn et al. 1995, Longán & Gómez-Bolea 1999), Navarre
(Etayo 2002) and Basque Country (Hawksworth & Etayo 2010) in the Iberian Peninsula. – 2, on
Flavoparmelia caperata on C. ladanifer, J. Etayo 29207.

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 317
*Acarospora epithallina H.Magn.
This species is a common member of saxicolous communities dominated by its host Acarospora hilaris
in southern and western Iberian Peninsula (Crespo et al. 1976). Reported from the area by Calatayud
et al. (2011a, b) where it is common (pers. obs.).
Acremonium subeffusum (Ellis & Galloway) Etayo & Brackel
This taxon was described as Dendrodochium subeffusum Ellis & Galloway growing on thalli of
Physcia millegrana but spreading also to Candelaria concolor (Brackel et al. 2012). Our sample was
found growing on dead Physconia venusta and fit well with the description of Dendrodochium subef-
fusum. It may represent the asexual phase of Pronectria leptaleae but the scarce number of records and
the lack of molecular data prevent confirmation of this relationship. – 2, on Physconia venusta on Q.
rotundifolia, J. Etayo 29075.
Actinocladium rhodosporum Ehrenb.
A common saprotrophic fungus growing on many different plant substrates (Ellis & Ellis 1985). It
is known from the Iberian Peninsula (Catalonia) growing on Rosmarinus officinalis (Muntañola-
Cvetkovic et al. 1997) and Buxus sempervirens (Muntañola-Cvetkovic et al. 2001). Brackel
(2013) found it growing on Flavoparmelia caperata in the Italian Alps. This specimen on Parmotrema
perlatum represents the second record growing as lichenicolous in the Iberian Peninsula (Etayo
2008a). – 2, on Parmotrema perlatum on branches of Q. rotundifolia, J. Etayo 29215.
*Agonimia tristicula (Nyl.) Zahlbr.
This species is commonly found growing on other lichens, especially on foliose saxicolous species
(Orange & Purvis 2009). – 2, on Physconia enteroxantha on acid rock, J. Etayo 29082.
Arthonia pepei Etayo & Pérez-Ortega, sp. nov. [MycoBank 817659] (Fig. 1)
Similar to Arthonia xanthoparmeliarum Etayo but lacking the characteristic necrotic black line sur-
rounding the infected area. The hymenium shows a bluish pigment absent in A. xanthoparmeliarum,
its paraphyses are not capitate and the ascospores are smaller.
Type: SPAIN, Extremadura, Cáceres, Monfragüe National Park, stands with Q. rotundifolia and C.
australis close to Monfragüe castle and hermitage, on Parmelina cryptotiliacea on Q. rotundifolia
branches, 13.04.2015, J. Etayo 29089 (holotype: MAF-Lich; isotype: hb. Etayo).
Description: Lichenicolous fungus on thallus of Parmelina cryptotiliacea, producing a pinkish to
cream coloured infection, roundish, 0.5 –1.2 mm diam., without black rim. Thalli with heavily infec-
ted areas turn completely bleached. Ascomata apothecioid, erumpent, soon sessile; firstly in small
number (1– 4), then forming small concentric circles up to 15 ascomata; rounded, flat to slightly con-
vex, 70 –140 µm diam., black, sometimes with a distinct very thin margin; when apothecia grow in
large numbers they form convex structures up to 370 µm in diam., with uneven borders. Exciple ab-
sent or much reduced, up to 20 µm thick, composed of hyphae slightly different from paraphysoids,
bluish-brown coloured, N+ orangish, K+ olivaceous sometimes with a black extracellular pigment.
Hymenium bluish grey, c. 30 µm high, I+ red, KI+ blue. Epihymenium slightly dark, almost black in
thick sections, N+ orangish, K+ olivaceous green. Paraphysoids abundant, immersed in a gel, simple
to branched, anastomosed, rarely septate, colourless in the central and lower parts, with thicker wall
and bluish brown pigment in the upper part, N+ orangish, K+ olivaceous, c. 2–3.5 µm wide and anticli-
nally or periclinally disposed regarding the asci. Subhymenium colourless, with some pigmented cells
with an extracellular, brown pigment, 10 –15 µm high. Asci semifissitunicate, clavate to ovate, stalked,
thickened apically, with apical beak, I+ red, KI+ blue, with a clear apical ring, 8-spored, 27–30 × 11–
13 µm. Ascospores colourless, 1-septate, ovoid or ovoid-fusiform, smooth, halonate, with upper cell
slightly wider, with one oil guttule in each cell, (8 –)9.5 –11 × 4 – 4.5 µm. Conidiomata not observed.
Etymology: Arthonia pepei is named after Josef Hafellner, who signed his correspondence to Spanish
lichenologists with Pepe, the Spanish hypocoristic of Josef. Together with Josef Poelt he was one of
the first central European lichenologists who regularly visited Spain in the seventies and eighties,

318 Herzogia 29 (2) Teil 1, 2016
Fig. 1: Arthonia pepei (holotype). A, B – Habitus on the thallus of Parmelina cryptotiliacea. C, E – Asci in KI. D –
Ascospore in water. – Scales: A–B = 100 µm; C–E = 10 µm.

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 319
contributing to the knowledge of Spanish lichen biota. Further, he has frequently helped many Spanish
lichenologists with puzzling species of lichen-forming and lichenicolous fungi.
Host and distribution: The new species grows on Parmelina cryptotiliacea, a recently described
species (Núñez-Zapata et al. 2011). It seems to be relatively abundant in Monfragüe National Park,
so we expect it also occurs in other parts of P. cryptotiliacea range.
Notes: Three species of Arthonia are known growing on the family Parmeliaceae (Lawrey &
Diederich 2015), namely Arthonia coronata Etayo on Flavoparmelia caperata, A. rimeliicola
Diederich on Parmotrema spp. and A. xanthoparmeliarum on Xanthoparmelia spp. Arthonia rimelii-
cola has much larger ascomata (0.15 – 0.4 mm in diam.) than A. pepei, which are covered by a white
and dense pruina, sometimes also with an orange to brown pruina in ascomatal rim. The ascospores are
(3 –)4-septate and much larger (16 –19 × 5 – 6.5 µm) than in A. pepei, and they show a clear perispore
becoming verrucose with age (Aptroot et al. 1997). Arthonia coronata is relatively common on so-
ralia of Flavoparmelia spp. although it has been recorded on Cladonia as well (Coppins & Aproot
2009). It is characterized by the convex, black, small sized (50 –100 µm) ascomata which are covered
by characteristic simple to septate setae (10 –35 × 3 –3.5 µm). Furthermore, hymenium and asci are
I–, KI– and ascospores are larger 11–14 × 3.5 –5 µm (Etayo 1996). The most similar taxon to A. pe-
pei is A. xanthoparmeliarum. It has similar sized ascomata 100 –150(–200) µm in diam. with whitish
dots in the center. However, the infections produced by A. xanthoparmeliarum are characteristically
surrounded by a black, necrotic rim. Furthermore, in this species the hymenium is colourless and the
epihymenium and hypothecium are brown, without the characteristic bluish pigment present in A. pe-
pei. In addition, paraphysoids in A. xanthoparmeliarum are typically capitate with large apical cells up
to 6 µm width and the ascospores are much larger (11–14.5 × 4 –5 µm) than in A. pepei, which further
show an amyloid perispore (Etayo & Sancho 2008). Arthonia muscigena Th.Fr. can also occur facul-
tatively on Parmeliaceae species (see hosts in Etayo & Sancho 2008). This taxon is lichenized and it
is easily distinguished by the thin thallus, the large ascomata (50 –250[–300] µm), the reddish brown
hypothecium, and the scarce and thin (0.5 –1 µm) paraphysoids (Coppins & Aptroot 2009).
Additional material studied: Arthonia coronata: Spain. Cantabria, Villaverde valley, Mollinedo, Monte Tejea,
track to Fuentes del Agüera, mixed forest; on Flavoparmelia caperata, 43º13'0.08''N, 3º16'15.5''W, 260 –350 m,
7.3.2014, J. Etayo 28504 (hb. Etayo). Arthonia xanthoparmeliarum: Chile. Magellanes Region, P.N. Pali-Aike,
on Xanthoparmelia sp. (sorediate) on volcanic boulder in the way to Morada del Diablo crater, 230 m, 2.2.2005, J.
Etayo 23111 (isotype).
Athelia arachnoidea (Berk.) Jülich
Plurivorous parasitic species, common in the Park growing on different hosts, even directly on mos-
ses. – 2, on bryophytes on C. australis, J. Etayo s.n.; ibidem, on thalli of Melanohalea exasperata,
Melanelixia subaurifera and Parmelina quercina, on Q. rotundifolia branches, J. Etayo 29208 (VIT).
4, on Physconia venusta on Q. rotundifolia, J. Etayo 29090.
Briancoppinsia cytospora (Vouaux) Diederich, Ertz, Lawrey & van den Boom
A generalistic parasitic lichenicolous fungus from the family Arthoniaceae growing on many different
hosts, very widespread (Diederich et al. 2012, Brackel 2014). It bleaches and necrotizes the infected
area, which does not show black rim around it. It is common in the Park growing on several species.
Melanelixia glabra and Physconia venusta represent new hosts for this species (Brackel 2014). – 2,
on Melanelixia glabra on Q. rotundifolia, J. Etayo 29076; ibidem, on Evernia prunastri on C. ladanifer,
J. Etayo 29078 (duplicate in VIT). 4, on M. glabra on Q. rotundifolia branches, J. Etayo 29083. – 3, on
Physconia venusta on F. angustifolia, J. Etayo 29097. 3. on M. glabra on F. angustifolia, J. Etayo 29104.
*Buellia badia (Fr.) A.Massal.
≡ Monerolechia badia (Fr.) Kalb
Common in the Iberian Peninsula (Llimona & Hladun 2001) and in the area of study. – 1, on
Protoparmeliopsis muralis on slates, J. Etayo 29144, 29145, 29147. – 5, on Xanthoparmelia tinctina
on quartzites, J. Etayo 29121. – 7, on Usnochroma carphinea on quartzites, J. Etayo 29131, 29133
(duplicate in VIT).

320 Herzogia 29 (2) Teil 1, 2016
Capronia triseptata (Diederich) Etayo
Known from many countries in Europe (Brackel 2014) including the Iberian Peninsula (Etayo 1996,
van den Boom & Etayo 2000). Also present in South America (Etayo 2002). This parasymbiont
(Diederich 1986) is a generalist able to grow both on crustose and foliose lichens. Previous Iberian
records were also found growing in the same host (Etayo 1996, van den Boom & Etayo 2000).
Perithecia are always covered by abundant (0 –1)2-septate setae in upper half. This character led
Etayo (1996) to combine this species from Muellerella into Capronia. Diederich (1986) pointed the
possibility of these setae being in fact a fungicolous hyphomycete. However the constant presence of
these setae in all samples and the constant absence of conidia make this possibility unlikely. – 2, on
apothecia of Parmelina quercina on Q. rotundifolia branches, J. Etayo 29209.
*Catillaria nigroclavata (Nyl.) Schuler
Common corticolous species in the Mediterranean Region (Llimona & Hladun 2001) frequenly oc-
curring on foliose lichens (Etayo 2010b). – 3, on Parmelina cryptotiliacea on Q. rotundifolia, J. Etayo
29093. – 6, on Ph. enteroxantha on Q. rotundifolia, J. Etayo 29137.
Cercidospora caudata Kernst.
A widespread taxon known from Europe, Africa (Morocco) and North America (Brackel 2014).
Some authors (e.g. Roux 2014) include this species within the concept of C. xanthoriae. So far, four
Cercidospora species are known growing on Teloschistaceae (C. caudata, C. epicallopisma Arnold,
C. epicarphinea [Nyl.] Grube & Hafellner and C. xanthoriae [Wedd.] R.Sant.). Our specimen fits well
in the concept of C. caudata although it differs in the 4-spored asci. The taxonomy of this group of
species is in need of a thorough revision. – 1, on Blastenia crenularia on slates, J. Etayo 29147.
Cercidospora lobothalliae Nav.-Ros. & Calat.
The species is also known from Greece, Spain, USA (California) and Russia (Navarro-Rosinés et al.
2009, Zhurbenko 2009). It grows on Lobothallia spp., mostly on L. radiosa although it has been also
found on L. melanaspis (Zhurbenko 2009). This is the first record on Circinaria. Cercidospora solea-
rispora Calat., Nav.-Ros. & Hafellner also grows on Circinaria caesiocinerea (Navarro-Rosinés et
al. 2009) but it is characterized by the larger ascomata and 1-septate (only rarely simple) ascospores. –
5, on Circinaria caesiocinerea on quartzitic outcrop, J. Etayo 29117.
Cercidospora macrospora (Uloth) Hafellner & Nav.-Ros.
Common species on apothecia of Protoparmeliopsis spp. Widespread in Europe, Asia, North America
and Africa (Calatayud et al. 2013). – 1, on Protoparmeliopsis muralis on slates, J. Etayo 29147.
Chalara sp.
Growing together with Lichenoconium erodens on brownish parts of Evernia prunastri thallus.
Unfortunately conidiogenous cells were old and conidia were not found. – 3, on Evernia prunastri on
F. angustifolia, J. Etayo 29101.
Cornutispora intermedia Punith. & D.Hawksw.
Widespread generalist, known from Germany (Brackel 2014), Italy (Brackel 2013), U.S.A.
(Punithalingam 2003) and Peru (Etayo 2010c). This is the first record on Ramalina (Brackel
2014) and apparently the first record for the Iberian Peninsula. – 3, on Ramalina fastigiata apothecia
on F. angustifolia, J. Etayo 29094.
Didymocyrtis cladoniicola (Diederich, Kocourk. & Etayo) Ertz & Diederich
Diederich et al. (2007) described this species growing on several Cladonia spp. It is widely distribu-
ted along Europe, and also occurs in North America, Africa (Canary Islands) and Asia (Hafellner
2015). In Monfragüe National Park it is commonly found growing on Cladonia foliacea squamules. –
5, on squamules of Cladonia foliacea, J. Etayo 29122.

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 321
Didymocyrtis ramalinae (Desm.) Ertz, Diederich & Hafellner
Ertz et al. (2015) established the teleomorph-anamorph relationship between this taxon and the re-
cently described Phoma ficuzzae Brackel (Brackel 2008b). Widely distributed along Europe and
known also from the Canary Islands, South Africa and Australia (Hafellner 2015). We observed
a phenotypic difference between specimens growing on the disk of Ramalina apothecia (perithecia
semiimmersed to almost sessile) and specimens growing on the thallus (perithecia completely im-
mersed). – 3, on Ramalina fastigiata on F. angustifolia, J. Etayo 29095.
*Diploschistes muscorum (Scop.) R.Sant.
Very abundant in the study area on several terricolous Cladonia species. – 5, on Cladonia spp. on quart-
zitic rock and soil, J. Etayo 29110; ibidem, on C. foliacea on quartzites, J. Etayo 29115, 29122, 29123.
Endococcus exerrans Nyl.
Several Endococcus species can be found growing on Rhizocarpon spp. Triebel (1989) reported
two species (E. perpusillus Nyl. and E. rugulosus (Borrer ex Leight.) Nyl.). Later, Sérusiaux et al.
(1999) established a narrower concept for the species growing on Rhizocarpon and provided morpho-
logical characters to separate three species. Etayo (2008b) further found E. macrosporus (Arnold)
Nyl. growing on R. geographicum. Endococcus species growing on Rhizocarpon urgently need a
thorough revision. Our specimen fits well with the concept of E. exerrans according to Sérusiaux
et al. (1999). Endococcus exerrans is known from Estonia (Suija & Randlane 2009), Germany
(Schiefelbein 2013), Italy (Brackel 2008a), Turkey (Halici 2008), Sweden (Thell et al. 2014),
Greece (Christensen & Alstrup 2013), La Gomera (Canary Islands) (van den Boom et al. 2015),
France (Roux 2012), the Netherlands (Aptroot et al. 2008) and Great Britain (Hawksworth 2003).
Roux & Egea (1992) reported the presence of a lichenicolous fungus in bad condition growing on
Opegrapha durieui Mont. on calcareous stones, which they identified as E. exerrans. Considering the
high host specificity shown by this species we consider this record belong to a different species. – 7,
on Rhizocarpon geographicum on quartzites, J. Etayo 29127.
Erythricium aurantiacum (Lasch) D.Hawksw. & A.Henrici
Generalist species, common on epiphytic lichens (especially from the Xanthorion community) along
Europe (Brackel 2014). – 2, on Physcia adscendens on branches of Q. rotundifolia, J. Etayo 29210.
Lichenochora weillii (Werner) Hafellner & R. Sant.
Cosmopolitan species, which grows on Physcia and Physconia spp. (Brackel 2014). It was found
forming small galls on Physconia thalli. The species was growing together with Reconditella physco-
niarum. – 1, on Physconia venusta on Q. rotundifolia, J. Etayo 29140, 29141.
Lichenoconium erodens M.S.Christ. & D.Hawksw.
Very common in Monfragüe, especially on Evernia prunastri thalli. – 2, on Evernia prunastri on C.
ladanifer J. Etayo 29078 (duplicate in VIT); ibidem, on thalli of Melanohalea exasperata on Q. rotun-
difolia branches, J. Etayo 29208 (duplicate in VIT). – 3, on E. prunastri on F. angustifolia, J. Etayo
29101. – 5, on E. prunastri, J. Etayo 29126.
Lichenoconium xanthoriae M.S.Christ.
Cosmopolitan, parasitic, generalist species growing on several host species. In Monfragüe it is parti-
cularly common occurring together with Diploschistes muscorum on Cladonia foliacea. Three other
Lichenoconium species [L. erodens, L. pyxidatae (Oudem.) Petr. & H.Sydow and L. usneae (Anzi) D.
Hawksw.] are known to occur on Cladonia but the size of conidia and conidiogenous cells fits with the
concept of L. xanthoriae. First record on Cladonia spp. Previously reported from the Iberian Peninsula
by van den Boom & Etayo (2000). – 5, on C. foliacea on quartzitic outcrops, J. Etayo 29115.
Lichenothelia rugosa (Thor) Ertz & Diederich
Very common species on Diploschistes spp. in the Iberian Peninsula. – 5, on Diploschistes scruposus
on quartzitic outcrop, J. Etayo 29109.

322 Herzogia 29 (2) Teil 1, 2016
Marchandiomyces corallinus (Roberge) Diederich & D.Hawksw.
Generalist species, grows on all kind of crustose, fruticose or foliose lichens (Brackel 2014). Known
from all continents except Australia and Antarctica (Brackel 2014). Very common growing on diffe-
rent epiphytic foliose hosts in the study area. – 2, on Xanthoparmelia tinctina on soil, J. Etayo 29071;
ibidem, on Melanelixia glabra on Q. rotundifolia, J. Etayo 29072; ibidem, on Physconia venusta on Q.
rotundifolia, J. Etayo 29073. – 4, on M. glabra on branches of Q. rotundifolia, J. Etayo 29083; ibidem,
on Physcia semipinnata and X. parietina on Q. rotundifolia branches, J. Etayo 29084; ibidem, on P.
cryptotiliacea on Q. rotundifolia branches, J. Etayo 29086. – 3, on Physcia adscendens on Q. rotun-
difolia twigs, J. Etayo 29103. – 5, on Xanthoparmelia tinctina on rock, J. Etayo 29112 (VIT); ibidem,
on E. prunastri, J. Etayo 29126.
Muellerella lichenicola (Sommerf.) D.Hawksw.
Cosmopolitan species, known from all continents. It occurs on many different host species, mostly on
crustose species, both saxicolous and epiphytic (Brackel 2014). – 3, on small apothecia of Caloplaca
sp. on Q. rotundifolia twigs, J. Etayo 29103.
Nesolechia fusca (Triebel & Rambold) Pérez-Ortega
Widespread taxon growing on several parmelioid hosts. Host specificity and the significance of asco-
spore size variability in Nesolechia are still in need of detailed molecular studies (Doré et al. 2006). –
5, on Xanthoparmelia tinctina on quartzites, J. Etayo 29116, 29121.
Niesslia cladoniicola D.Hawksw. & W.Gams
Hawksworth (1975) described ascospore size as 8 –10.5 × 1.5 –2.2 µm, although several authors
have repeatedly reported smaller sizes (Etayo & Sancho 2008, Zhurbenko & Alstrup 2004). On
the other hand, our specimen growing on Cladonia foliacea has slightly larger ascospores (8 –13 ×
2.5 –3 µm). Thus, it is not clear whether several taxa are currently included in the concept of N. cla-
doniicola or ascospore size varies more widely than originally proposed. Recently reported from the
Iberian Peninsula by van den Boom & Etayo 2014. – 5, on squamules of C. foliacea, J. Etayo 29123.
Paranectria oropensis (Cesati in Rabenh.) D.Hawksw.
Plurivorous species, known from several host species, mostly in epiphytic communities where this
taxon plays an important role in community dynamics (Hafellner & Obermayer 2009). We found
it abundantly, either alone or growing together with Trichonectria rubefaciens on Parmelina crypto-
tiliacea. – 3, on P. cryptotiliacea on F. angustifolia, J. Etayo 29096. – 5, on P. cryptotiliacea on Q.
rotundifolia, J. Etayo 29106.
Phaeospora everniae Etayo & van den Boom
Recently described from the Iberian Peninsula (van den Boom & Etayo 2014). It has also been recor-
ded recently from France (Roux et al. 2016) and U.K. (http://www.britishlichensociety.org.uk/resour-
ces/lichen-taxon-database/taxon-detail/21365). – 3, on thallus of Evernia prunastri on F. angustifolia,
J. Etayo 29101. – 6, on soralia of E. prunastri on Q. rotundifolia, J. Etayo 29138.
*Polysporina subfuscescens (Nyl.) K.Knudsen & Kocourk.
Lichenicolous species of the genus Polysporina have been recently reviewed by Knudsen &
Kocourková (2008). These authors showed that the name Polysporina lapponica had been mis-
applied to lichenicolous specimens. The type of this species is lichenized and belongs to the genus
Sarcogyne (Knudsen & Kocourková 2008). Subsequently, lichenicolous specimens previously
adscribed to P. lapponica were identified as P. subfuscescens. Frequent in the study area growing on
several saxicolous host species. –1, on Blastenia crenularia on slates, J. Etayo 29145. – 5, on several
crustose lichens on quartzites, J. Etayo 29119.
Pronectria leptaleae (J.Steiner) Lowen
= Xenonectriella leptaleae (J.Steiner) Rossman & Lowen

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 323
Rossman et al. (1999) combined this species into the genus Xenonectriella in spite of the K+ pale
brown to black reaction of the perithecial wall. Our specimens either do not react with K or present
a K brown reaction. Thus, we think it is better to include it in the genus Pronectria. This species is
rather abundant in Monfragüe National Park on thallus and apothecia of Physconia spp. Our specimens
further differ from the description provided by Rossman et al. (1999) in the verruculose ascospores
(pale golden-brown and prominently tuberculate with age in Rossman et al. 1999). This species is usu-
ally found together with Acremonium subeffusum, so they may represent different states of the same
taxon. – 2, on Physconia venusta on Q. rotundifolia, J. Etayo 29074. – 3, on P. venusta on F. angustifo-
lia, J. Etayo 29097. – 5, on P. venusta and P. enteroxantha on Q. rotundifolia, J. Etayo 29113. – 4, on P.
venusta on Q. rotundifolia, J. Etayo 29087. – 6, on P. enteroxantha on Q. rotundifolia, J. Etayo 29135.
Pronectria subimperspicua (Speg.) Lowen
Considered to be a tropical species, it has been recently recorded from Germany on Parmelia sul-
cata, Physcia tenella and Punctelia jeckeri (Brackel 2014). We found it growing together with the
common Paranectria oropensis. First record for the Iberian Peninsula. –5, on P. cryptotiliacea on Q.
rotundifolia, J. Etayo 29106, 29107.
Pronectria tincta (Fuckel) Lowen
= P. santessonii (Lowen & D.Hawksw.) Lowen
Pronectria santessonii was introduced by Lowen & Hawksworth (1986) for specimens growing
on Anaptychia runcinata. These authors also noted the presence of an undescribed third species of
Pronectria growing on Anaptychia. Etayo (2010b) discussed the variability of ascospore shape, size
and ornamentation of Pronectria specimens growing on Anaptychia (and rarely on Physcia spp.) and
proposed the synonymy of P. santessonii with P. tincta based on the overlap in ascospore size and
shape between species. – 2, on Anaptychia ciliaris on branches of Q. rotundifolia, J. Etayo 29211.
Reconditella physconiarum Hafellner & Matzer
Holarctic species. It is rather common growing on Physconia spp. in Monfragüe, and more rarely on
Phaeophyscia orbicularis, which represents a new host for this species. – 1, on Physconia venusta on
Q. rotundifolia, J. Etayo 29140. – 2, on Physconia enteroxantha on acid stone, J. Etayo 29082; ibidem,
on Physconia sp. on branches of Q. rotundifolia, J. Etayo 29214. –5, on P. enteroxantha (rarely) on
Q. rotundifolia, J. Etayo 29113. – 6, on P. enteroxantha on Q. rotundifolia, J. Etayo 29137. – 7, on
Phaeophyscia orbicularis on P. nigra, J. Etayo 29127.
Roselliniella cladoniae (Anzi) Matzer & Hafellner
Widespread taxon on several Cladonia spp. Although it has been previously recorded from La Gomera
in Spain (Hafellner 1996), this is the first record for the Iberian Peninsula. – 3, on Cladonia sp.
squamules on rock, J. Etayo 29099. – 5, on Cladonia sp. squamules on rock, J. Etayo 29111.
Sphaerellothecium reticulatum (Zopf) Etayo
A common species on different hosts of the family Parmeliaceae (Brackel 2014). This is the first re-
cord on Evernia prunastri. – 1, on Xanthoparmelia tinctina on slates, J. Etayo 29146. – 2, on Evernia
prunastri on C. ladanifer, J. Etayo 29081. – 5, on Evernia on Q. rotundifolia, J. Etayo 29126, ibidem,
on Xanthoparmelia tinctina on quartzites, J. Etayo 29121. – 6, on E. prunastri on Q. rotundifolia, J.
Etayo 29139.
Stigmidium squamariae (de Lesd.) Roux & Triebel
Widespread on Protoparmeliopsis, Rhizoplaca and lobate Lecanora species in the Northern Hemisphere
(Brackel 2014). – 1, on apothecia of Protoparmeliopsis muralis on slates, J. Etayo 29144, 29145, 29150.
Syzygospora physciacearum Diederich
Cosmopolitan on several foliose species of Physciaceae, common in the Iberian Peninsula (Llimona
& Hladun 2001). – 2, on Physcia adscendens on leaves of B. sempervirens, J. Etayo 29213. – 3, on
P. adscendens on twigs of an unidentified shrub, J. Etayo 29103.

324 Herzogia 29 (2) Teil 1, 2016
Taeniolella phaeophysciae D.Hawksw.
Holarctic species, which grows on various genera of Physciaceae, common in the Iberian Peninsula
(Llimona & Hladun 2001). In Monfragüe we found it growing on Phaeophyscia orbicularis and
Physconia spp. together with Reconditella physconiarum. – 2, on thalli of Physconia sp. on Q. rotun-
difolia branches, J. Etayo 29208 (VIT); ibidem, on Physconia sp. on branches of Q. rotundifolia, J.
Etayo 29214. – 7, on Phaeophyscia orbicularis on P. nigra, J. Etayo 29127.
Tremella caloplacae (Zahlbr.) Diederich
Millanes et al. (2014) provided a complete distribution of this species. In the Iberian Peninsula this
species is known growing on Usnochroma carphinea (Sérusiaux et al. 2003) and on Blastenia ceri-
na, Athallia holocarpa, Xanthocarpia marmorata and Variospora velana (Etayo 2010b) but not on
Xanthoria. – 3, on Xanthoria parietina on Q. rotundifolia twigs, J. Etayo 29092. – 7, on X. parietina
on P. nigra, J. Etayo 29128.
Tremella cladoniae Diederich & M.S.Christ.
We found this species growing abundantly on Cladonia foliacea, mostly on podetia and on apothecia.
It has been widely reported from many countries in Europe (Diederich 1996) but not previously from
the Iberian Peninsula. – 5, on Cladonia foliacea on quartzitic rocks, J. Etayo 29105.
Trichonectria rubefaciens (Ellis & Everh.) Diederich & Schroers
A relatively common species in the Iberian Peninsula (van den Boom & Etayo 2000, Etayo 2002,
Etayo & López de Silanes 2008). Parmelina cryptotiliacea and Evernia prunastri represent new
hosts for this species, indicating a plurivorous behavior on Parmeliaceae. Common in Monfragüe
National Park. Perithecia usually grow surrounded by Acremonium rhabdosporum W.Gams, its pu-
tative anamorph (Lowen 1995). Our specimens fit well within the concept of A. rhabdosporum alt-
hough we did not observed the annular thickenings of the wall at the apices pointed by Hawksworth
(1979). – 2, on Parmelina cryptotiliacea on Q. rotundifolia, J. Etayo 29080. – 4, on P. cryptotiliacea
on Q. rotundifolia branches, J. Etayo 29085; ibidem, J. Etayo 29086. – 3, on P. cryptotiliacea on Q.
rotundifolia branches, J. Etayo 29091; ibidem, on Evernia prunastri on Fraxinus, J. Etayo 29100. – 5,
on E. prunastri (Acremonium state), J. Etayo 29126.
Unguiculariopsis lettaui (Grummann) Coppins
A very common species in the Iberian Peninsula, but not be frequent in Monfragüe. – 3, on Evernia
prunastri on F. angustifolia, J. Etayo 29101.
Xanthoriicola physciae (Kalchbr.) D.Hawksw.
Cosmopolitan species, growing on X. parietina and more rarely on X. polycarpa (Brackel 2014). It has
been recently shown that together with other lichenicolous fungi it belongs to the Teratosphaeriaceae
(Ruibal et al. 2011, Pérez-Ortega et al. 2015). – 2, on apothecia of Xanthoria parietina on Q. rotun-
difolia branches, J. Etayo 29208 (VIT).
Zwackhiomyces echinulatus Brackel
Three Zwackhiomyces species are known to grow on hosts of the family Physciaceae. Zwackhiomyces
physciicola Alstrup (Alstrup 1993) was described from Denmark on Physcia caesia. It has rela-
tively thin-walled asci (78 –92 × 11.5 –13 µm) and smooth-walled, non-halonate ascospores, (18 –22 ×
5.5 – 6.5 µm). Spanish material of this species on P. caesia always has ascospores with three pairs of setu-
lae, which may indicate that it belongs to a different genus (Sérusiaux et al. 1999, Etayo 2010). Brackel
(2008a) described Z. echinulatus on Physconia distorta. This taxon is characterized by the small pe-
rithecia (100 –250 µm), 4(6 – 8)-spored asci (82–100 × 13 –17 µm), and ascospores (24 –27 × 9.3 –11 µm)
with echinulate perispore. Kocakaya et al. (2011) described Z. turcicus Kocakaya, Halici & Aksoy on
Physcia magnussonii. It is characterized by the large perithecia (200 – 450 µm), the (4 –)6(– 8)-spored
asci (70 –92 × 19 –21 µm) and by the ascospores with verruculose ornamentation (18 –22 × 8 –9.5 µm).
Our specimen on Physconia venusta mostly fits the concept of Z. echinulatus. Perithecia are rather small,
asci are either 4 or 6-spored (68 – 85 × 14 –17 µm) and ascospores are similar in size (20 –25 × 8 –11 µm).

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 325
However, ascospores show a verrucose perispore rather than echinulate. First record for the Iberian
Peninsula. – 1, on Physconia venusta on Q. rotundifolia, J. Etayo 29149 (hb. Etayo).
Acknowledgements
We thank to P. Diederich (Luxembourg) for his help with Tremella cladoniae,W. Sanders (Ft. Myers) for improving
the English and E. Ros (Pamplona) for accompanying the first author to Monfragüe N.P. SPO is supported by the grant
RYC-2014-16784 from the Spanish Economy and Competitiveness Ministry.
References
Alstrup, V. 1993. News on lichens and lichenicolous fungi from the Nordic countries. – Graphis Scripta 5: 96 –104.
Aptroot, A., Sérusiaux, E. & Sipman H. J. M. 1997. Lichens and Lichenicolous Fungi from New Guinea. –
Bibliotheca Lichenologica 64: 1–220.
Aptroot, A., Brand, M., van Herk, K. & Sparrius, L. 2008. Veranderingen in de checklist van de Nederlandse
korstmossen en korstmosparasieten. – Buxbaumiella 82: 6 –13.
Atienza, V., Pérez-Ortega, S. & Etayo, J. 2009. Two new conidial lichenicolous fungi from Spain indicate the
distinction of Lichenodiplis and Minutoexcipula. – The Lichenologist 41: 223 –229.
Brackel, W. v. 2008a. Zwackhiomyces echinulatus sp. nov. and other lichenicolous fungi from Sicily, Italy. –
Herzogia 21: 181–198.
Brackel, W. v. 2008b. Phoma ficuzzae sp. nov. and some other lichenicolous fungi from Sicily, Italy. – Sauteria 15:
103 –120.
Brackel, W. v. 2013. Miscellaneous records of lichenicolous fungi from the Italian Alps. – Herzogia 26: 141–157.
Brackel, W. v. 2014. Kommentierter Katalog der flechtenbewohnenden Pilze Bayerns. – Bibliotheca Lichenologica
109: 1– 476.
Brackel, W. v., Etayo, J. & Lechat, C. 2012. Notes on three lichenicolous species of Acremonium including two
new species. – Mycosphere 3: 854 – 862.
Calatayud, V. & Etayo, J. 2001. Five new species of lichenicolous conidial fungi from Spain. – Canadian Journal
of Botany 79: 223 –230.
Calatayud, V., Corrales, J. M. & Hernández, S. 2011a. Guía de los líquenes del Parque Nacional de Monfragüe. –
Cáceres: Universidad de Extremadura, Servicio de Publicaciones.
Calatayud, V., Corrales, J. M. & Hernández, S. 2011b. Líquenes del Parque Nacional de Monfragüe. – Cáceres:
Universidad de Extremadura, Servicio de Publicaciones.
Calatayud, V., Navarro-Rosinés, P. & Hafellner, J. 2013. Contributions to a revision of Cercidospora
(Dothideales), 2: Species on Lecanora s.l., Rhizoplaca and Squamarina. – Mycosphere 4: 539 –557.
Christensen, S. N. & Alstrup, V. 2013. Notes on epilithic, epigeic and muscicolous lichens and lichenicolous fungi
from rock outcrops in the mountains of northern Greece. – Mycobiota 1: 25 –50.
Coppins, B. J. & Aptroot, A. 2009. Arthonia Ach. (1806). – In: Smith, C. W., Aptroot, A., Coppins, B. J.,
Fletcher, A., Gilbert, O. L., James, P. W. & Wolseley, P. A. (eds). The Lichens of Great Britain and Ireland.
Pp. 153 –171. – London: British Lichen Society.
Crespo, A., Barreno, E. & Follmann, G. 1976. Sobre las comunidades líquénicas rupícolas de Acarospora hilaris
(Duf.) Hue en la Península Ibérica. – Anales del Instituto Botanico A. J. Cavanilles 33: 189 –205.
Diederich, P. 1986. Lichenicolous fungi from the Grand Dutchy of Luxembourg and surrounding areas. – Lejeunia
119: 1–26.
Diederich, P. 1996. The lichenicolous heterobasidiomycetes. – Bibliotheca Lichenologica 61: 1–198.
Diederich, P., Kocourková, J., Etayo, J. & Zhurbenko, M. 2007. The lichenicolous Phoma species (Coelomycetes)
on Cladonia. – The Lichenologist 39: 153 –163.
Diederich, P., Lawrey, J. D., Sikaroodi, M., van den Boom, P. P. & Ertz, D. 2012. Briancoppinsia, a new coelo-
mycetous genus of Arthoniaceae (Arthoniales) for the lichenicolous Phoma cytospora, with a key to this and
similar taxa. – Fungal Diversity 52: 1–12.
Doré, C. J., Cole, M. S. & Hawksworth, D. L. 2006. Preliminary statistical studies of the infraspecific variation in
the ascospores of Nesolechia oxyspora growing on different genera of parmelioid lichens. – The Lichenologist
38: 425 – 434.
Ellis, M. B. & Ellis, J. P. 1985. Microfungi on land plants. An identification handbook. – London: Croom Helm Ltd.
Ertz, D., Diederich, P., Lawrey, J. D., Berger, F., Freebury, C. E., Coppins, B., Gardiennet, A. & Hafellner,
J. 2015. Phylogenetic insights resolve Dacampiaceae (Pleosporales) as polyphyletic: Didymocyrtis (Pleosporales,
Phaeosphaeriaceae) with Phoma-like anamorphs resurrected and segregated from Polycoccum (Trypetheliales,
Polycoccaceae fam. nov.). – Fungal Diversity 74: 53 – 89.

326 Herzogia 29 (2) Teil 1, 2016
Etayo, J. 1996. Aportaciόn a la flora liquénica de las Islas Canarias. I. Hongos liquenícolas de Gomera. – Bulletin de
la Societè Linnéenne de Provence 47: 93 –110.
Etayo, J. 2002. Catálogo de líquenes y hongos liquenícolas del Parque Natural de Bértiz (Navarra, España). – Bulletin
de la Societè Linnéenne de Provence 53: 155 –170.
Etayo, J. 2008a. Líquenes y hongos liquenícolas del LIC de Ablitas (S. Navarra, España). – Cryptogamie Mycologie
29: 63 –94.
Etayo, J. 2008b. Líquenes y hongos liquenícolas de Aiko-Harria. – http://www.lifeaiakoharria.net/datos/documentos/
Estudio%20de%20 los%20l%C3 %ADquenes%20de%20AH_2008.pdf
Etayo, J. 2010a. Lichenicolous fungi from the western Pyrenees. V. Three new ascomycetes. – Opuscula Philolichenum
8: 131–139.
Etayo, J. 2010b. Líquenes y hongos liquenícolas de Aragón. – Guineana 16: 1– 501.
Etayo, J. 2010c. Hongos liquenícolas de Perú. Homenaje a Rolf Santesson. – Bulletin de la Societè Linnéenne de
Provence 61: 1– 46.
Etayo, J. & Sancho, L. G. 2008. Hongos liquenícola del Sur de Sudamérica, especialmente de Isla Navarino
(Chile). – Bibliotheca Lichenologica 98: 1–302.
Etayo, J. & López de Silanes, M. E. 2008. Líquenes epífitos y hongos liquenícolas del Bosque Viejo de Munain-
Okariz (Álava, País Vasco, España). – Nova Acta Científica Compostelana. Bioloxía 17: 11–29.
Glenn, M. G., Gomez-Bolea, A. & Lobello, R. 1995. Metal content and community structure of cryptogam bioin-
dicators in relation to vehicular traffic in Montseny Biosphere Reserve (Catalonia, Spain). – The Lichenologist
27: 291–304.
Hafellner, J. & Casares-Porcel, M. 2003. Lichenicolous fungi invading lichens on gypsum soils in southern
Spain. – Herzogia 16: 123 –133.
Hafellner, J. 1996. Bemerkenswerte Funde von Flechten und lichenicole Pilze auf Macaronesischen Inseln IV.
Einige bisher übersehene lichenicole Arten der Kanarischen Inseln. – Cryptogamie, Bryologie, Lichénologie 17:
1–14.
Hafellner, J. 2015. Distributional and other data for some species of Didymocyrtis (Dothideomycetes, Pleosporales,
Phaeosphaeriaceae), including their Phoma-type anamorphs. – Fritschiana 80: 43 – 88.
Hafellner, J. & Obermayer, W. 2009. The role of Paranectria oropensis in community dynamics of epiphyte syn-
usia on roadside trees. – Herzogia 22: 177–190.
Halici, M. G. 2008. Key to the lichenicolous Ascomycota (including mitosporic fungi) of Turkey. – Mycotaxon 104:
253 –286.
Hawksworth, D. L. 1975. Notes on British lichenicolous fungi, I. – Kew Bulletin 30: 183 –203.
Hawksworth, D. L. 1979. The lichenicolous hyphomycetes. – Bulletin of the British Museum (Natural History).
Botany (Bot.) 6: 183 –300.
Hawksworth, D. L. 2003. The lichenicolous fungi of Great Britain and Ireland: an overview and annotated check-
list. – The Lichenologist 35: 191–232.
Hawksworth, D. L. & Etayo, J. 2010. Calongia gibelluloides gen. sp. nov., a remarkable minute new lichenicolous
conidial fungus from northern Spain. – The Lichenologist 42: 355 –359.
Knudsen, K. & Kocourková, J. 2008. A study of lichenicolous species of Polysporina (Acarosporaceae). –
Mycotaxon 105: 149 –164.
Kocakaya, M., Halıcı, M. G. & Aksoy, A. 2011. Zwackhiomyces turcicus sp. nov. (Ascomycota, Xanthopyreniaceae)
from Turkey. – Mycotaxon 116: 329 –333.
Lawrey, J. D. & Diederich, P. 2015. Lichenicolous fungi – worldwide checklist, including isolated cultures and
sequences available. – http://www.lichenicolous.net [10/10/2015].
Llimona, X. & Hladun, N. L. 2001. Checklist of the lichens and lichenicolous fungi of the Iberian Peninsula and
Balearic Islands. – Bocconea 14: 1–581.
Llop, E. & Gómez-Bolea, A. 2006. Foliicolous lichens and associated lichenicolous fungi in the north-eastern
Iberian Peninsula: the effect of environmental factors on distribution. – The Lichenologist 38: 55 – 65.
Longán, A. & Gómez-Bolea, A. 1999. Líquenes y hongos liquenícolas epífitos de Quercus ilex L., poco conocidos
en encinares de Cataluña (España). – Cryptogamie Mycologie 20: 49 –55.
Lowen, R. & Hawksworth, D. L. 1986. Nectriella santessonii, a new lichenicolous pyrenomycete with an
Acremonium anamorph. – The Lichenologist 18: 321–328.
Lowen, R. 1995. Acremonium section Lichenoidea section nov. and Pronectria oligospora species nov. – Mycotaxon
53: 81–95.
Martínez, I. 2002. Lichenicolous fungi from the Iberian Peninsula and the Macaronesian area. – Nova Hedwigia
74: 51– 67.
Millanes, A., Diederich, P., Westberg, M., Knutsson, T. & Wedin, M. 2014. Tremella rhizocarpicola sp. nov.
and other interesting lichenicolous Tremellales and Filobasidiales in the Nordic countries. – MycoKeys 8:
31– 41.

Etayo & Pérez-Ortega: Lichenicolous fungi and lichens from Monfragüe National Park 327
Muntañola-Cvetkovic, M. & Navarro, P. H. 1997. Més fongs mitospòrics de les terres baixes del sur i de l‘oest
de Catalunya (Espanya). – Revista Catalana de Micologia 20: 63 – 83.
Muntañola-Cvetkovic, M., Hoyo, M., Sierra D. & Llimona, X. 2001. Lesions i estructures fúngiques en arbus-
tos espontanis de Catalunya. III. Branques i troncs de Buxus sempervirens. – Revista Catalana de Micologia 24:
17– 41.
Navarro-Rosinés, P., Hladun, N. L. & Llimona, X. 2008. Gelatinopsis heppiae sp. nov. (Leotiales, Ascomycota)
un hongo liquenícola sobre Heppia despraeuxii de Aragón (NE de España). – Revista Catalana de Micologia 30:
57– 64.
Navarro-Rosinés, P., Calatayud, V. & Hafellner, J. 2009. Contributions to a revision of the genus Cercidospora
(Dothideales) 1. Species on Megasporaceae. – Mycotaxon 110: 5 –25.
Núñez-Zapata, J., Divakar, P. K., Ruth, D. P., Cubas, P., Hawksworth, D. L. & Crespo, A. 2011. Conundrums
in species concepts: the discovery of a new cryptic species segregated from Parmelina tiliacea (Ascomycota:
Parmeliaceae). – The Lichenologist 43: 603 – 616.
Orange, A. & Purvis, O. W. 2009. Agonimia Zahlbr. (1909). – In: Smith, C. W., Aptroot, A., Coppins, B. J.,
Fletcher, A., Gilbert, O. L., James, P. W. & Wolseley, P. A. (eds). The Lichens of Great Britain and Ireland.
Pp. 136 –138. – London: British Lichen Society.
Pérez-Ortega, S. & Etayo, J. 2008. A new species of Lecanora s. lat., growing on Lasallia pustulata. – The
Lichenologist 40: 111–118.
Pérez-Ortega, S. & Álvarez-Lafuente, A. 2006. Primer catálogo de líquenes y hongos liquenícolas de la
Comunidad Autónoma de Castilla y León (España). – Botanica Complutensis 30: 17–52.
Pérez-Ortega, S., Garrido-Benavent, I. & De Los Ríos, A. 2015. Austrostigmidium, a new austral genus of
lichenicolous fungi close to rock-inhabiting meristematic fungi in Teratosphaeriaceae. – The Lichenologist 47:
143 –156.
Punithalingam, E. 2003. Nuclei, micronuclei and appendages in tri-and tetraradiate conidia of Cornutispora and
four other coelomycete genera. –Mycological Research 107: 917–948.
Rivas-Martínez, S. 1987. Memoria del Mapa de Series de Vegetación de España. – Madrid: ICONA.
Rossman, A. Y., Samuels, G. J., Rogerson, C. T. & Lowen, R. 1999. Genera of Bionectriaceae, Hypocreaceae and
Nectriaceae (Hypocreales, Ascomycetes). – Studies in Mycology 42: 1–248.
Roux C. et coll. 2016. Liste des lichens et champignons lichénicoles de France métropolitaine (mise à jour
2016/01/25). – http://lichenologue.org/fr/.
Roux, C. 2012. Liste des lichens et champignons lichénicoles de France. – Bulletin de la Societè Linnéen de Provence
16: 1–220.
Roux, C. 2014. Catalogue des lichens et champignons lichénicoles de France métropolitaine. – Fougeres: Henry des
Abbayes.
Roux, C., & Egea, J. M. 1992. L’Opegraphetum durieui Egea et Roux ass. nov., une association lichénique saxicole-
calcicole, halophile. – Cryptogamie Bryologie Lichénologie 13: 105 –115.
Ruibal, C., Millanes, A. M. & Hawksworth, D. L. 2011. Molecular phylogenetic studies on the lichenicolous
Xanthoriicola physciae reveal Antarctic rock-inhabiting fungi and Piedraia species among closest relatives in the
Teratosphaeriaceae. – IMA Fungus 2: 97–103.
Schiefelbein, U. 2013. Additions to the lichenized and lichenicolous fungi of Mecklenburg-Western Pomerania
(Germany). – Herzogia 26: 65 –72.
Sérusiaux, E., Diederich, P., Brand, A. M. & van den Boom, P. 1999. New or interesting lichens and lichenicolous
fungi from Belgium and Luxembourg. VIII. – Lejeunia 162: 1–95.
Sérusiaux, E., Diederich, P., Ertz, D. & van den Boom, P. P. G. 2003. New or interesting lichens and lichenicolous
fungi from Belgium and Luxembourg. IX. – Lejeunia 173: 1– 48.
Suija, A. & Randlane, T. 2009. Excluded species of lichenized and lichenicolous fungi. – Folia Cryptogamica
Estonica 46: 87– 88.
Thell, A., Alstrup, V., Arup, U., Bendiksby, M., Czarnota, P. et al. 2014. New or interesting lichens and licheni-
colous fungi from the Vadstena area, Östergötland, Sweden. – Graphis Scripta 26: 15 –33.
Triebel, D. 1989. Lecideicole Ascomyceten. Eine Revision der obligat lichenicolen Ascomyceten auf lecideoiden
Flechten. – Bibliotheca Lichenologica 35: 1–278.
van den Boom, P. P. G. & Etayo, J. 2000. Contribution to the knowledge of lichenicolous fungi and lichens from
Portugal and Spain. – Österreichische Zeitschrift für Pilzkunde 9: 151–162.
van den Boom, P. P. G. & Etayo, J. 2014. New records of lichenicolous fungi and lichenicolous lichens from the
Iberian Peninsula, with the description of four new species and one new genus. – Opuscula Philolichenum 13:
44 –79.
van den Boom, P. P. G., Clerc, P. & Ertz, D. 2015. New Records of lichens and lichenicolous fungi from
La Gomera (Canary Islands, Spain), including the new species: Usnea boomiana P. Clerc. – Candollea 70:
165 –177.

328 Herzogia 29 (2) Teil 1, 2016
Zamora, J. C., Pérez-Ortega, S. & Rico, V. J. 2011. Tremella macrobasidiata (Basidiomycota, Tremellales), a new
lichenicolous fungus from the Iberian Peninsula. – The Lichenologist 43: 407– 415.
Zhurbenko, M. P. 2009. Lichenicolous fungi and some lichens from the Holarctic. – Opuscula Philolichenum 6:
87–120.
Zhurbenko, M. P. & Alstrup, V. 2004. Lichenicolous fungi on Cladonia mainly from the Arctic. – Symbolae
Botanicae Upsalienses 34: 477– 499.
Manuscript accepted: 30 July 2016.
Communicated by: Lucia Muggia
Addresses of the authors
Javier Etayo, Navarro Villoslada, 16-3° dcha., 31003 Pamplona, Spain.
E-mail: jetayosa@educacion.navarra.es
Sergio Perez-Ortega, Real Jardín Botánico (CSIC), Plaza Murillo 2, 28014, Madrid, Spain.
E-mail: sperezortega@rjb.csic.es