The lichen genus Porina in Macaronesia, with descriptions ... · The lichen genus Porina in...

The Lichenologist 39(1): 15–33 (2007) � 2007 British Lichen Societydoi:10.1017/S0024282907005993 Printed in the United Kingdom

The lichen genus Porina in Macaronesia, with descriptions oftwo new species

Emmanuël SÉRUSIAUX, Franz BERGER, Maarten BRAND andPieter van den BOOM

Abstract: Detailed studies on the lichen genus Porina in Macaronesia have led to a reappraisal of thegenuine identity of Porina atlantica (Erichsen) P. M. Jørg., a characteristic species that has previouslybeen confused with P. guaranitica, P. heterospora, P. nucula, P. mastoidea or P. rhodostoma in theliterature, and is here reported from Madeira, the Canary Islands, Ireland, France and Portugal. Twonew species are described: P. effilata Brand & Sérus. sp. nov. (known from Madeira, the CanaryIslands, Great Britain, Ireland and Portugal) and P. ocoteae Brand & Sérus. sp. nov. (restricted to LaPalma, Canary Islands, and São Jorge in the Azores). Porina leptospora Nyl. is recognized at the specieslevel, and P. isidiata Kalb & Hafellner is reduced into synonymy with P. atlantica. A key to all knownspecies of Porina in Macaronesia is provided.

Key words: Azores, Canary Islands, England, France, Gomera, Ireland, La Palma, laurisilva,Los Tilos, Madeira, Porina effilata, Porina ocoteae, Portugal, rare species, São Jorge, Tenerife, Wales

Introduction

The checklist of lichens occurring inMacaronesia (Hafellner 1995, 1999, 2002)reflects the confusion of authors with regardto the identity of several species of Porina inthe archipelagos. For example, Hafellner(1995: 74) suspected that earlier reports ofP. mastoidea (Ach.) A. Massal. and P. nuculaAch. referred to P. isidiata Kalb & Hafellner;he accepted P. guaranitica Malme as adistinct species, with P. heterospora (Fink)R. C. Harris as a synonym. For the samegroup of species, Etayo (1998: 100–101)and McCarthy (2003: 21) accepted only onespecies, viz. P. atlantica (Erichsen) P. M.Jørg., with P. guaranitica and P. heterosporaas synonyms.

As we now have large collections of Porinafrom Madeira and the Canary Islands as wellas some from the Azores, we are able toproduce a more detailed taxonomic accountof the species present. The large amountof additional relevant data now available(Harris 1995; Lücking & Vezda 1998;McCarthy, 1993, 2000, 2001, 2003) wasalso critical for this study.

The generic concept

Almost simultaneously, Harris (1995) andKalb & Hafellner (1995) introduced newgeneric concepts within the Porinaceae (thevalid name for the family since the typifica-tion of Porina Müll. Arg. with P. nuculaAch.; see McCarthy 2003: 8), includingrevisions of the circumscription of the well-known genus Porina. The two taxonomiccircumscriptions are not consistent witheach other and neither has received widesupport (see McCarthy & Malcolm 1997 fora discussion), although the genus Pseudo-sagedia (Müll. Arg.) M. Choisy, resurrectedby Kalb & Hafellner (1995), is now used inseveral European checklists. The purpose of

E. Sérusiaux: Plant Taxonomy & Conservation BiologyUnit, Environmental Sciences Dept., University ofLiège, Sart Tilman B22, B-4000 Liège, Belgium.Email: [email protected]. Berger: A-4794 Kopfing 130, Austria.A. M. Brand: Klipperwerf 5, NL-2317 DX Leiden, TheNetherlands.P. van den Boom: Arafura 16, NL-5691 JA Son, TheNetherlands.

this paper is to examine the species presentin Macaronesia, and, to some extent, thosealso present in Western Europe and isthus species-dedicated. We have thereforeadopted a conservative generic conceptand broad delimitation of Porina, followingMcCarthy (2003). We have also suggestedthat the introduction of new generic con-cepts should take into account phylogeneticanalyses based on multiple gene DNAsequences, when representative species of allgroups within the family have been tested(Grube et al. 2004a; Baloch & Grube,2006).

MethodsThis study is based on collections and observationsmade by the authors in Macaronesia, western Europeand several tropical regions, as well as on the examina-tion of herbarium material borrowed from institutionsand colleagues. The material was examined in tapwater, in lactophenol cotton-blue (LCB; FLUKAChemika 61335) or in Lugol’s solution (IKI; Lugolsolution SIGMA L-6146). The measurements alwaysrefer to material mounted in water. The dimensions ofperithecia are those of the perithecia or when presentthe perithecial warts (enclosing the perithecia s. str.) inherbarium specimens.

The Species

Porina atlantica (Erichsen)P. M. Jørg.

Graphis Scripta 12: 1 (2000).—Ocellularia atlanticaErichsen, Hedwigia 66: 276 (1926); type: CanaryIslands, Tenerife, Llano de los Vieijos in Monte Minasde Abajo, on Laurus canariensis, 17 iii 1919, L. Lindiger(HBG—holotypus, not seen) [material requested butnot received].

Porina isidiata Kalb & Hafellner, Herzogia 9: 83(1992); type: Madeira, An der Levada da Serra do Faialentlang, etwas SW von Santo da Serra in RichtungCamacha, in letzten Resten eines alten, jedoch starkgestörten, NW-exponierten Eucalyptus-Waldes, anLaurus azorica, 750 m, 13 viii 1990, K. Kalb 23484 (hbKalb—holotypus!, isotypus!).

Porina guaranitica auct. europ. p. p., non Malme,Ark. Bot. 23(A): 13 (1929).

Porina heterospora auct. europ. p. p., non (Fink) R. C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980).

Porina mastoidea auct. europ. p. p., non (Ach.) Müll.Arg., Bot. Jahrb. Syst. 6: 399 (1885).

Porina nucula auct. europ. p. p., non Ach., Syn. Meth.Lich.: 112 (1814).

Porina rhodostoma auct. europ., non Müll. Arg., Bot.Jahrb. Syst. 6: 398 (1885).

(Figs 1A & B, 3A–D, 7A)

Thallus epiphloeodal, rather thick(c. 0·5 mm) and strongly appressed andadherent to the bark, occasionally overgrow-ing liverworts and mosses, very rare onrocks, usually not shiny, dark green to palegreyish, in herbarium specimens usually palewith an orange or pinkish tinge, when freshand alive typically orange-brown to vividgreen, rugose to verrucose, sometimesirregularly wrinkled, typically with oxalatecrystals deposited in globose agglomerationsinside the thallus (crystallocumuli sensuHafellner & Kalb 1995: 163–164), whichcan usually be seen as translucent dots underhigh magnification, without a prothallus.Isidia absent or locally developed on parts ofthalli, or abundant and covering large partsof thalli, simple to coralloid, cylindrical tomost commonly globose and constrictedseveral times along their length and thusappearing moniliform, usually brittle andeasily removed, concolorous with the thallusor paler, 0·1–0·5(–1·0) mm high.

Perithecia rare to abundant, hemisphericalto subglobose, slightly but distinctly con-stricted at their base, rarely not, 0·5–0·9(–1·1) mm diam., typically covered by athallus layer containing agglomerations ofoxalate crystals and thus appearing as ifenclosed in a large thalline wart; ostiolarregion slightly depressed, usually orange,pinkish, reddish to brownish, rarely darkbrown, in young perithecia almost indistinctas the covering thallus layer has not yet gotthinner so that the underlying orange involu-crellum becomes visible. Involucrellum typi-cally pale orange in section and K+ brightorange to reddish, 30–40 �m thick, readilydetached from the thallus covering layer insections and separated from it by a very loosehyphal tissue around the base of matureperithecia. Excipulum pale yellow, c. 20 �mthick. Paraphyses abundant, simple or rarelybranched near the base, not inflated at theapices, c. 1·5 �m thick. Asci cylindrical-clavate, up to 200�20–24 �m. Ascospores8/ascus, fusiform, never tapering towardsone end, straight or slightly curved, 7–9(–13)-septate (one spore seen with 15

16 THE LICHENOLOGIST Vol. 39

septa), 44–80�7–13 �m, plus a distinct,2–3 �m thick perispore.

Pycnidia rare, usually immersed in the thal-lus with only the ostioles visible, 0·1–0·3 mm

diam., with a hyaline or pale orange, K+bright orange wall. Pycnidiospores ellipsoid,sometimes slightly tapering towards one end,hyaline, simple, 2·5–3·0�1·0–1·5 �m.

F. 1. Morphology of Porina species. A & B, P. atlantica [La Palma, Barranca Gallegos, iv 2004, E. Sérusiaux s.n. (LG)]; C & D, P. effilata, arrows point to perithecia and colours electronically exaggerated in C to make theperithecia more conspicuous [C: Ireland, Killarney, 2000, A. M. Brand 40242 (hb Brand); D: La Palma, Cubo dela Galga, 3 iv 1986, A. M. Brand 13302 (hb Brand)]; E & F, P. ocoteae, E, young perithecia; F, mature perithecia

[La Palma, Los Tilos, iv 2004, E. Sérusiaux s. n. (LG—holotypus)]. Scale=0·1 mm.

2007 Porina in Macaronesia—Sérusiaux et al. 17

Notes. In Macaronesia, Porina atlantica is adistinctive species, easily recognized by itsdull, verrucose thallus containing packets ofoxalate crystals, large perithecia covered bya thallus layer with an orange to reddishbrown periostiolar region, and 7–9(–13)-septate ascospores, 44–80�7–13 �m, nottapering towards one end and with a thickperispore. Quite surprisingly, it has beenconfused with P. effilata, newly described inthis paper, and which has a non-verrucosethallus (lacking packets of oxalate crystals),smaller perithecia with a smooth or slightlytomentose surface, and ascospores typicallyeffilate (caudate and tapering towards theproximal end), narrower [9–11(–13)-septate, 68–83�7–11 �m] and with a muchthinner perispore. Porina atlantica comesclose to P. nucula Ach., characterized by athallus that is strongly appressed and adher-

ent to the bark, verrucose, pale orange tovivid green when fresh and without a proth-allus, prominent perithecia covered by athallus layer with a slightly reddish ostiolarregion and broad ascospores, 7-septate andwith a thick perispore.

Porina nucula Ach. (Figs 2C, 3E & F) isthe type species of the genus, but several,often confused taxa are clearly related. Theirtaxonomy remains to be clarified, as demon-strated by the preliminary treatment ofHarris (1995: 172–175) for Florida (USA).The lectotypification made by Harris (1995:174) points to a species with peritheciaslightly constricted at their base andwith ascospores 40–52�11–14 �m, withrounded ends. He further demonstrated aconsiderable variation in ascospore size andseptation in the same group, under theinformal ‘‘sp. 674’’ [ascospores narrower,

F. 2. Morphology of Porina species. A, P. guaranitica [Paraguay, 1893, G. O. A. Malme 1469 (S—holotypus)];B, P. mastoidea [Papua New Guinea, Laing Island, 1992, P. Diederich 11644 (hb Diederich)]; C, P. nucula [PapuaNew Guinea, Burbura logging site, 28 vii 1992, P. Diederich 11909b (hb Diederich)]; D, P. rhodostoma [Dominica,

Cabrit National Park, 26 iii 1998, E. Sérusiaux s. n. (LG)]. Scale=0·1 mm.


with pointed ends, 42–53(–61)�(7–)8–11(–12) �m], ‘‘sp. 6’’ [ascospores withends pointed, �evenly tapering at bothends, 60–75�10–13(–15) �m; perithecialwarts large, c. 0·6 mm in diam.], and ‘‘sp.30308’’ (ascospores 25–33�5–6 �m;perithecial warts small, c. 0·3 mm in diam.).The matter is indeed unresolved, and ourown experience with tropical material con-firms that a great deal of variation is presentand that further taxa are most probablyinvolved. We therefore decided to adoptthe ‘‘finely cut’’ circumscription of Harris

(1995: 174) and Lücking & Vezda (1998:213–214). R. Lücking kindly provided uswith data on the lectotypes designated byR. C. Harris in H and NY. The delimitationof the species by McCarthy (1994: 400;2001: 140) is broader and would questionthe circumscription adopted here for P.atlantica. Until more data become availablefrom tropical areas, we distinguish these twospecies mainly by the size of the peri-thecia: in P. altantica, as circumscribed here,mature perithecia are 0·5–0·9(–1·1) mm indiameter, while they never exceed 0·7 mm inP. nucula.

Within the populations here assigned to P.atlantica, septation and size of ascosporescan vary considerably: several specimensfrom Macaronesia and Ireland (Fig. 3A & B)consistently have 7-septate ascosporesmeasuring 44–60�7–11 �m, while in othersthe spores are more septate and longer. Themost divergent specimens are ‘Madeira,Chão da Ribeira, vii 2003, E. Sérusiaux s. n.’which has 9–13-septate ascospores measur-ing 60–71�10–13 �m (Fig. 3C), and‘SW France, Ste Engrâce, P. van denBoom 12821’ which has 9–11(–15)-septateascospores measuring 72–80�11–13 �m(Fig. 3D). As the variation covers the wholedistributional range (as studied in thispaper) of the species and is not correlatedwith any other characters, it has beenincluded in the general description providedabove.

Examination of the species names formerlyused for the species in the area of study. Tavares(1952: 320) used the name ‘‘Porina mastoi-dea (Ach.) Mass.’’ for specimens of thisspecies collected in Madeira, although hehad not seen the type. However, he didexamine the type of Ocellularia atlanticaErichsen, described from Tenerife, and hewas ‘‘inclined to think that the Tenerifelichen is only a variant of Porina mastoidea’’.The description provided by Erichsen(1926: 276) reads as follows: ‘‘Thallus (. . .)verruculoso-inaequalis, interdum aliquidrimosus, (. . .). Apothecia solitaria in verru-cis. Verrucae fertiles subglobosae, basi nonconstrictae, numerosae sed non confertae,

F. 3. Ascospores of Porina species. A–D, P. atlantica[A, Ireland, West Cork, 20 ix 1982, P. W. James s. n.(BM); B, La Palma, Los Tilos, iv 2004, E. Sérusiaux s.n. (LG); C, Madeira, Chão da Ribeira, vii 2003, E.Sérusiaux s. n. (LG); D, France, Ste Engrâce, P. van denBoom 12821 (hb van den Boom)]; E & F, P. nucula[E, Papua New Guinea, P. Diederich 11489 (LG, hbDiederich); F, Papua New Guinea, P. Diederich 11909b

(LG, hb Diederich)]. Scale:=10 �m.


0·6–1 mm latae. (. . .) Sporae octonae,decoloratae, fusiformes, 5–11 septatae,0·062–70 mm longae et 0·018–21 mmcrassae, episporio 0·003–4 mm crasso.’’ It isa rather accurate description of the taxondealt with here. Although the type collectionwas not available for this study, we acceptthis epithet.

Porina mastoidea (Ach.) Müll. Arg. is adifferent species with a rather smooth, shiny,thick thallus which readily peels andbecomes detached from the substratum,and forms bullate portions, with a well-developed black prothallus. Its perithecia areusually applanate and rarely constricted attheir base, typically have a conspicuousblack ostiole, and the ascospores are7-septate, 32–66�6–13 �m, with a 1·5–3·5 �m thick perispore (ascospore charactersfollowing McCarthy 2001: 137; Figs 2B, 4A

& B). Although the ascospores are quitesimilar to those of P. nucula and P. atlantica,P. mastoidea probably belongs to a differenttaxonomic group, which corresponds toClathroporina Müll. Arg. sensu Harris (1995:171–172). Porina africana Müll. Arg., P.tetracerae (Ach.) Müll. Arg. and P. ocoteaeBrand & Sérus., newly described in thispaper, also belong to that latter group.

No material of P. atlantica was available toSwinscow when he published his outstand-ing revision of the genus in the British Isles(Swinscow 1962): indeed, both collectionshe mentioned under P. nucula Ach. fromIreland belong to P. effilata (pp. 47–49; bothcollections seen in BM: Aug. 1935, D. A.Jones s. n. and June 26, 1961, P. W. James &T. D. V. Swinscow s. n.; see below, under P.effilata). The drawings of ascospores (withinthe ascus or standing alone) in Swinscow’sfig. 12 are typical for P. effilata, but hissection of a perithecium is misleading as wellas part of his description of the perithecialwall: indeed, he records that perithecia are‘‘largely covered by a thalline layer’’ andadds the puzzling ‘‘loculi containing plaquesof refractile material sometimes present inthalline covering layer’’. This is never thecase with P. effilata nor in the material fromIreland he had at hand. We suggest thatSwinscow had extracted these data on theperithecia from other specimens he hadexamined, apparently from tropical areasand belonging to the genuine P. nuculaor other species, possibly the genuine P.heterospora [see his sentence (p. 48): ‘‘Thedescription is based on the examination offive specimens from the southern UnitedStates of America, two from Cuba, and threefrom south-west Ireland’’]. He may haveseen caudate and multi-septate ascosporesfrom material belonging to P. heterosporafrom the US (Fig. 4C–E) and these areindeed close to the Irish material of P. effilatahe had on his table. The drawing of theperithecial section in Swinscow’s fig. 12 is afurther clue to the confusion of species as theascospores it contains are not at all caudateand mostly 7-septate, and thus cannotbelong to either of the Irish specimenshe was studying. This mixture of data is

F. 4. Ascospores of Porina species. A & B, P. mastoi-dea [A, Papua New Guinea, P. Diederich 11491 (LG, hbDiederich); B, Papua New Guinea, P. Diederich 11644(LG, hb Diederich)]; C–E, P. heterospora [C, USA,Florida, E. Sérusiaux 1765 (LG); D, USA, Louisiana,S. C. Tucker 18481 (E); E, Guatemala, P. van den Boom

33491 (LG, hb van den Boom)]. Scale=10 �m.


suspected to be the source of the confusionfor further workers examining material of P.atlantica and P. effilata.

Swinscow (1962: 47–49) assigned the twocollections from Ireland he was studying toP. nucula Ach., a widespread pantropicalspecies (McCarthy 2003: 73–74). The dif-ferentiation of this species from P. atlanticais quite difficult and has been discussedabove, the main conspicuous differencebeing the size of mature perithecia.

The use of the epithet nucula was acceptedby Poelt & Vezda (1977: 195, 201) for‘‘Ireland and Portugal’’ and by Champion(1976: 30) for a collection from ‘‘Anaga’’ onTenerife in the Canary Islands. Both P.atlantica and P. effilata are now recognizedin Ireland and Portugal, while only P.atlantica is known in the Sierra de Anaga inTenerife.

Following the suggestion by Harris (1975:173), the name P. heterospora (Fink) R. C.Harris (=‘‘P. nucula auct. brit., non Ach.’’)was adopted in the The Lichen Flora of GreatBritain and Ireland (Purvis et al. 1992: 491).However, it is clear from the descriptionprovided that the authors of the Flora wereworking with material of both P. atlanticaand P. effilata. Indeed, meanwhile P. atlan-tica had been found in SW Ireland (collec-tions made in 1982 by P. W. James andP. M. Jørgensen) and its conspicuous andlarge perithecia were noted by theselichenologists. The description of thallusand perithecia published in Purvis et al.(1992) reads as follows and was clearlyinspired by Swinscow’s drawing (1962: 47):‘‘Thallus (. . .) coarsely granular to verru-cose, sometimes �corrugate-wrinkled.Perithecia 0·5–0·9 mm diam., (. . .) largelycovered by a thalline exciple containinglocules with plaque-like crystals, immersedin thalline warts with only apex exposed’’; itperfectly matches P. atlantica. The descrip-tion of ascospores (‘‘Ascospores 40–80�8–12 �m, 7–14-septate, fusiform-clavate’’)refers to P. effilata and was also inspired bySwinscow’s drawing.

The epithet heterospora was introduced inthe lichen flora of Macaronesia by Gonzálezet al. (1990: 106) for Tenerife and by Kalb &

Hafellner (1992: 82) for Madeira. The areastudied by González et al. is a part of theSierra de Anaga where P. atlantica occurs,and the species is also common in all laurelforests of Madeira.

Kalb & Hafellner (1992: 83) described P.isidiata Kalb & Hafellner as the isidiatecounterpart of specimens they referred to P.heterospora but are obviously identical withP. atlantica. We suggest that P. isidiata ismerely a form of P. atlantica of no taxonomicvalue. Isidiate specimens of P. atlantica havebeen observed in all localities sampledin Macaronesia, as well as in Europeanmaterial. The isidia are globose to cylindricaland sometimes form coralloid proliferationson the thallus surface; they can be sparseand present only on parts of the thalli andseem to develop into coralloid outgrowthsin specimens under stress. The distinctionbetween abnormal outgrowths of the photo-biont and genuine isidia in Porina is a diffi-cult matter: the experience of McCarthy(1993: 16) suggests ‘‘these structures aremanifestations of rampant growth by thephotobiont’’, an opinion shared by Harris(1995: 171). However, the latter authordescribed the new Clathroporina isidiifera onthe basis for example of its ‘‘true isidia,constricted at the base with an organizationsimilar to that of the thallus’’. It can indeedbe suggested that two different types of‘‘isidia’’ can be found in the genus, onebeing genuine isidia and acting as diaspores,and the other being ‘‘adaptations to enhancegas exchange in a permanently humidenvironment’’ (Grube et al. 2004b: 1161). Asimilar situation is found within populationsof another species found mainly in La Palma(P. ocoteae; see below): many individuals donot produce such isidioid outgrowths, butthose in apparently stressed conditions orgrowing in less suitable habitats do producethem; moreover, such thalli do not produceperithecia. In these cases (isidioid thalli in P.atlantica and P. ocoteae), we are convincedthat the production of ‘‘isidia’’ does notrepresent a diagnostic criterion for taxo-nomic purposes. Otherwise, the collectionsreferred to P. isidiata are identical with P.atlantica.


Jørgensen (2000a) examined the type ofOcellularia atlantica Erichsen and found it tobe identical with the populations named P.heterospora ‘‘distributed from SW Ireland toMacaronesia’’. The photograph provided ofthe type clearly indicates that it representsthe species dealt with here as P. atlantica.However, the only collection made byJørgensen in SW Ireland and with a hand-written identification label as P. atlanticaseen by us (P. M. Jørgensen no. 9121, BG!)is typical P. effilata. He also examined(Jørgensen 2000b) an isidiate specimen healso had collected in SW Ireland during thesame field trip in 1982 (P. M. Jørgensen no.9126, BG!) and accepted the epithet isidiatafor it. This specimen has a few isidia anddefinitely belongs to the isidiate form of P.atlantica.

In his revision of the saxicolous Porinain the Southern Hemisphere, McCarthy(1993: 49) reduced P. heterospora (Fink) R.C. Harris into synonymy with P. guaraniticaMalme, described from rocks in Paraguay.This opinion was adopted by van den Boomet al. (1995: 278) for a collection from SWFrance, but was questioned by Harris (1995:173). McCarthy (2003: 21) confirmed thisview in his remarkable Catalogue of thePorinaceae, and further accepted the olderepithet atlantica for the species. We believethat McCarthy adopted a broad species con-cept for this group, which includes P. guara-nitica from S America, the P. heterosporagroup as studied in detail by Harris (1995:172–175) for the south-eastern UnitedStates, P. atlantica as circumscribed hereand the newly described P. effilata.

The type collection of P. guaranitica hasbeen examined (Paraguay, Paraguaría, onbasalt, 27 vi 1893, G. O. A. Malme 1469,S—lectotype). It grows on rock, its peri-thecia (immersed in thalline verrucae; seeMcCarthy 1993: 50, fig. 50) do not exceed0·5 mm in diam. and have a pale orangeperiostiolar region; its ascospores areelongate-fusiform, slightly but distinctlytapering towards one end, 10–16-septate,74–87�9–10 �m (plus a perispore 2 �mthick; Fig. 2A, 5A). Harris (1995: 173) alsosaw the type but gave different data on spore

septation and size: .9·5 septaZ and 57–71 �m long for ten spores examined. Thespecimens named P. atlantica here usuallyhave larger perithecia with a larger anddarker periostiolar region, and ascosporesusually with fewer septa, and never taperingtowards one end. Although we have notstudied any other specimen from SouthAmerica, we suggest that these differencesmight be worth recognition at the speciesrank. However, even if the populations fromSouth America are to be considered as con-specific with those from Macaronesia, atlan-tica is the older epithet and can be used forthe populations found in Macaronesia andWestern Europe.

In his study of the Trichotheliaceae fromFlorida (USA), Harris (1995: 172–175)described P. heterospora (Fink) R. C. Harriswith clavate spores having a long tapering

F. 5. Ascospores of Porina species. A, P. guaranitica[Paraguay, G. O. A. Malme 1469 (S—lectotype)]; B &C, P. rhodostoma [B, Cuba, C. Wright, (G—holotype);C, Dominica, 26 iii 1998, E. Sérusiaux s. n. (LG)]; D,P. leptospora [Azores, F. Berger 15974 (hb Berger)]; E &F, P. borreri [E, France, Ste Engrâce, 10 vii 1989, E.Sérusiaux s. n. (LG); F, Belgium, Furfooz, P. van den

Boom 13835 (LG)]. Scale=10 �m.


tail, (9–)11–13-septate, 85–110�12–14 �m. We can confirm such a descriptionon the basis of specimens from SE USA andGuatemala (see list below; Fig. 4C–E). Suchshape, septation and size are completelydifferent from either species dealt with here(P. atlantica or P. effilata) and the epithetheterospora can thus be excluded from theflora of Europe and Macaronesia. Porinaheterospora is said (Harris 1995: 173) to becommon in the southern coastal plain of SEUSA, as well as Cuba and Venezuela.

McCarthy & Malcolm (1996: 549) intro-duced the epithet ‘‘aff. rubrostoma Müll.Arg.’’ (a mispelling of rhodostoma Müll.Arg.) for a collection from Tenerife (LasMontanas de Anaga). The correspondingspecimen has not been examined but weassume it belongs to P. atlantica, as twospecimens have been seen from the verysame area in Tenerife. McCarthy (2003: 84)has also used the epithet rhodostoma for thespecimen from SW Ireland named isidiata byJørgensen (2000b), which we here refer toP. atlantica (see above). The epithet nowappears in the Checklist of Great Britain andIreland (Coppins 2002: 47; see also http://users.argonet.co.uk/users/jmgray/checklist.html, visited on Dec. 29th, 2004,no longer in use and replaced by http://www.thebls.org.uk/checklist.html#P, visitedon July 31th, 2006). Besides the type col-lection and these other two specimens, P.rhodostoma has never been cited elsewhere inthe literature (McCarthy 2003: 84).

The type collection of P. rhodostoma wasalso examined (Cuba, C. Wright ‘‘Ser. II,539, G—holotype). It grows on bark, has arugulose to verrucose thallus, large andrather flattened perithecia immersed inthallus verrucae (up to 0·8 mm diam.), witha conspicuous, large, typically dark red(dark red to red-brown) periostiolar region.The original description states that theascospores are ‘‘fusiformes, 65–80 �mlongae et 10–12 �m latae, 7–11-septatae,loculi 2 intermedii reliquis longiores’’. Twoperithecia have been sectioned and theascospores found outside the asci are ratherbadly preserved; they are largely fusiform,with rounded ends, not tapering towards

one end, 5–9-septate, 41–64�13–17 �m(plus a 3 �m thick perispore; Fig. 5B). Wehave seen no collection from Macaronesiathat could match such characteristics.

A healthy, recently collected specimenfrom Dominica, West Indies (Fig. 2D; seebelow), is referred to the same species;its thallus and perithecia are identical, itsascospores have the same shape but are11–13-septate and measure 74–89�21–26 �m (plus a 2–3 �m thick perispore;Fig. 5C). If further collections can confirmthese observations, it would mean that theascospores studied in the type are not yetfully mature. We suggest that these twocollections represent a different taxon fromP. atlantica, differing mainly by the typicallydark red and large periostiolar region and bythe size and septation of the ascospores.

Ecology and distribution. Porina atlanticaoccurs in all stands of the laurisilva studiedin Madeira (where it can be common) andthe Canary Islands; so far, no specimenshave been collected in the Azores. It usuallygrows on bark in rather shaded places andhas been recorded on basalt boulders inLa Palma, inside well-preserved laurisilvawhere the species is rather common on trees.It is also known from several localitiesalong the Atlantic coast of western Europewhere it seems to be very rare: SW Irelandin two localities, France (Brittany and thewestern Pyrenees), and Portugal (Sintra). InBrittany, the thallus of the only collectionavailable is typical (isidioid outgrowths arealso present), but no mature peritheciacould be seen. It must also be noted that allEuropean localities are well-known for theirwell-developed lichen flora comprisingseveral typical atlantic species.

Specimens examined. France: Dépt. Finistère: 26 kmW of Huelgoat, forêt de Cranou, near St-Conval, on oldQuercus, 175 m, 1996, A. M. Brand 34439 (hb Brand).Dépt. Pyrénées Atlantiques: Ste Engrâce, near entrance ofGorges of Kakouetta, shaded wood, on Buxus, 510 m,1992, P. van den Boom 12821 (hb van den Boom).—Ireland: V. C. H2, North Kerry: Cromaglown bridge,on Quercus, 1982, P. M. Jørgensen 9126 (BG—L/48004,mentioned as P. isidiata by Jørgensen 2000b); Killarney,Meeting of the Waters, N of old Weir Bridge, on oldsheltered Quercus at shore, 2000, A. M. Brand 40241


(hb Brand). V. C. H3, West Cork: Glengarriff woods,shaded side of old Quercus, 20 ix 1982, P. W. Jamess. n. (BM).—Portugal: W of Lisboa, Sintra, nearCapuchos monastery, on old Cupressus, 250 m, 2003,A. M. Brand 49770 (hb Brand).—Canary Islands:Gomera: Parque Nacional de Garajonay, El Cedro,chemin depuis le village jusqu’à l’Ermita, le long dubarranco del Cedro, laurisylve de fond de vallée, avecPersea indica, 900–950 m, 26 vii 1994, E. Sérusiaux s. n.(LG, 2 collections). La Palma: 8·5 km N of Santa Cruz,Bco la Galga, Cubo de la Galga, laurisilva forest in cleft,550 m, 1999, P. van den Boom 22430 (hb van denBoom); ibid., on basalt boulder inside the forest, 580 m,1986, A. M. Brand 13313 (hb Brand); Los Tilos (W deLas Lomados), laurisilve riche en Hedera canariensiset en fougères (dont Woodwardia radicans), on bark,800–850 m, vii 1997, E. Sérusiaux s. n. (LG); ibid.,600–700 m, iv 2004, E. Sérusiaux s. n. (LG); ibid., nearpicnic place, on old Octoea, 450 m, 1986, A. M. Brand13558 (hb Brand); Barranco Gallegos, croisementde la route Barlovento-Roque Faro par la laguna deBarlovento, rochers de basalte en sous-bois, 900 m, iv2004, E. Sérusiaux s. n. (LG). Tenerife: Laurisilva, headof San Andres Valley near El Bailadero, Sierra deAnaga, 1964, H. A. & F. H. Imshaug 35802 (LG);Las Mercedes, on trees, 600–700 m, xii 1984, C.Hdez Padrón (E).—Madeira: Ribeiro Frio, Hänge SWoberhalb der Fischzuchtanstalt, in Lorbeerwald miteinzelnen Kastanienbäumen, aus Laurus azorica,950 m, 1990, K. & A. Kalb 23710 (hb Kalb); RibeiroFrio, le long de la Levada de Portela, laurisylve�dégradée, 850 m, ii 1988, E. Sérusiaux s. n. (LG); S deSeixal, Chão da Ribeira, début du chemin montant versFanal, laurisylve peu perturbée, 500 m, vii 2003, E.Sérusiaux s. n. (LG); Rabaçal, entre les maisons deRabaçal et la Cascata do Risco, fourrés d’Erica enbordure de la levada, 1050 m, vii 2003, E. Sérusiauxs. n. (LG); Fanal (le long de la route Ribeira de Janelavers Paul da Serra), vieux Ocotea foetens dans un pâtur-age, 1100 m, vii 2003, E. Sérusiaux s. n. (LG); RouteRibeira Brava/São Vicente, un peu au N de Boca deEncumeada, fourrés du Fayal-Brezal avec quelquesfûts, 900 m, vii 2003, E. Sérusiaux s. n. (LG); Casa dasQueimadas, promenade vers Caldeirão Verde, fourrésd’Erica en bordure de la levada et arbres plantés, 900 m,vii 2003, E. Sérusiaux s. n. (LG).

Selected specimens examined for other species mentioned.Porina heterospora (Fink) R. C. Harris: USA: Florida:

along road no. 361 between Adams Beach and thehighway ALT27, Spring Warrio creek, on Cupressusby a swamp, 5 m, 1976, E. Sérusiaux 1765 (LG).Louisiana: East Feliciana Parish, Idlewild ExperimentalFarm, c. 3 miles SE of Clinton, 30(48#N 90(45#W, onQuercus in downed hardwoods, 1979, S. C. Tucker18481 (E).—Guatemala: Dept. Alta Verapaz: NW ofCoban, ‘‘Las Victorias’’ forest in the national park,15(28.4#N 90(22.8#W, on trunk of small tree, 1320 m,2004, P. van den Boom 33491 (hb van den Boom, LG).

Porina nucula Ach.: Papua New Guinea: Madangprov.: near Bogia, along road Bogia-Josephstaal, 4(27#S144(56#E, on trunk of felled trees among gardens,

330 m, 1992, P. Diederich 11489 (LG, hb Diederich);Burbura logging site, c. 30 km NNW of Madang,4(50#S 145(38#E, on tree in virgin rainforest on lowhills, 70 m, 1992, P. Diederich 11901 & 11909b (LG, hbDiederich).

Porina mastoidea (Ach.) Müll. Arg.: Papua NewGuinea: Madang prov.: near Bogia, along road Bogia–Josephstaal, 4(27#S 144(56#E, on trunk of felled treesamong gardens, 330 m, 1992, P. Diederich 11491 (LG,hb Diederich); near Bogia, Laing Island in Hansa Bay,4(10#S 144(52#E, corticolous in coastal forest oncoral island, 1 m, 1992, P. Diederich 11644 (LG, hbDiederich).

Porina rhodostoma Müll. Arg.: West Indies:Dominica: Cabrit National Park, disturbed dry forest,on tree, c. 100 m, 26 iii 1998, E. Sérusiaux s. n. (LG).

Porina effilata Brand & Sérus. sp. nov.

A speciebus generis in Europa et Macaronesia differt asporis caudatibus, (7–)9–11(–13)-septatis, (65–)68–83(–85)�7–11 �m.

Typus: Canary Islands, Gomera, Parque Nacional deGarajonay, El Cedro, chemin depuis le village jusqu’àl’Ermita, le long du barranco del Cedro, laurisilve defond de vallée, avec Persea indica, sur tronc, 900–950 m,26 July 1994, E. Sérusiaux s. n. (LG—holotypus).

Porina atlantica auct. europ. p. p., non (Erichsen)P. M. Jørg., Graphis Scripta 12: 1 (2000).

Porina guaranitica auct. europ. p. p., non Malme,Ark. Bot. 23(A): 13 (1929).

Porina heterospora auct. europ. p. p., non (Fink) R. C.Harris, in Tucker & Harris, Bryologist 83: 12 (1980).

Porina nucula auct. europ. p. p., non Ach., Syn. Meth.Lich.: 112 (1814).

(Figs 1C & D, 6A–C, 7B)

Thallus mainly epiphloeodal, able to over-grow corticolous liverworts and mosses,crustose and rather thin, green to dark greenor pale beige and �pinkish, sometimes witha tinge of blue, rather matt, rarely shiny, notcontaining large oxalate crystals, without aprothallus. Isidia very rare (seen only onparts of thalli in a few collections fromIreland), simple, almost globose, c. 0·1 mmhigh, not glossy and usually almost powdery,easily detached from the thallus.

Perithecia rare or abundant, at firstimmersed in the thallus and occasionallyremaining so for a long time (and thereforehardly visible), or protuberant and sessile,not included in thallus-dominated verrucae,sometimes remaining hidden in cracks of thebark or under bryophytes, subglobose, 0·3–0·45(–0·5) mm diam.; perithecia initially


covered by a thin, greenish thallus layer thatcan remain for a long time around the base,pale orange to orange, sometimes pinkish,rarely almost translucent, K+ bright orangeto deep red, sometimes distinctly and finelygreyish-whitish tomentose in the upper half,rarely slightly verruculose, with a centralostiole appearing as a pale or translucent dotor as a hole. Involucrellum well-developed,especially in upper parts, separated from theexcipulum in old and overmature perithecia,especially near the base, c. 20–25 �m thick.Excipulum pale orange, c. 15–20 �m thick.Paraphyses usually abundant, simple orrarely branched near the base, not inflated atthe apices, 1–1·5 �m thick. Asci cylindrical-clavate, up to 180�20 �m when fullymature. Ascospores 8/ascus, narrowly

fusiform, slightly but distinctly inflated inthe upper half and tapering towards theproximal end, straight or usually slightlycurved, (7–)9–11(–13)-septate, (65–)68–83(–85)�7–11 �m, with a thin but distinctperispore.

Pycnidia very rare, �immersed in thethallus with only the ostioles visible, 0·1–0·2 mm diam., with a pale orange, K+bright orange wall. Pycnidiospores cylindrical,straight or slightly curved, with roundedends, hyaline, simple, 15–17(–18)�c. 1 �m.

Notes. This new species is easily dis-tinguished by its pale orange, sometimespinkish, and rather small perithecia, notincluded in thallus warts (albeit sometimesremaining immersed in the thallus and thushardly visible, or sometimes covered by athin thallus layer in young stages that canremain around their base), sometimes with adistinct tomentum in their upper half, andby its long, distinctly tapering, mainly 9–11-septate ascospores. Its thallus never containsagglomerations of large oxalate crystals. Itdoes not belong to the same group as P.atlantica with which it has been confused; wesuggest it comes close or even belongs toSegestria Fr. sensu Harris (1995: 175–176) orto the Porina rufula group sensu Lücking(2004: 410, 416–417).

No close relative has been found in theliterature. Porina sylvatica McCarthy &Kantvilas, known from SE Australia(McCarthy & Kantvilas 1993: 144–145),has larger (0·43–0·72 mm diam.), smoothand orange-brown to reddish brown peri-thecia and its ascospores are narrower (6·5–9·5 �m), not tapering towards their proximalends. Porina speciosa McCarthy & Malcolm,from New Zealand (McCarthy & Malcolm1996), has larger (0·56–1·01 mm diam.),smooth perithecia and much largerascospores (65–122�15–22 �m) with athick perispore.

As explained above under P. atlantica,Porina effilata has been mentioned in theEuropean literature under many differentepithets: P. nucula (Swinscow 1962: 47–49),P. heterospora (Purvis et al. 1992: 491), P.guaranitica (McCarthy 1993: 49–54), and P.

F. 6. Ascospores of Porina species. A–C, P. effilata[A, Gomera, El Cedro, 26 vii 1994, E. Sérusiaux s. n.(LG—holotypus); B, La Palma, Bco la Galga, A. M.Brand 13302 (hb Brand); C, Ireland, West Cork,Glengarriff, 11 viii 1966, P. W. James s. n. (BM)]; D &E, P. ocoteae [D, La Palma, Los Tilos, iv 2004, E.Sérusiaux s. n. (LG—holotypus); E, La Palma, LosTilos, P. van den Boom 22174 (hb van den Boom)]; F,P. africana [Guatemala, P. van den Boom 33423 (hb vanden Boom, LG)]; G, P. tetracerae [Rwanda, 30 iii 2005,

E. Sérusiaux s. n. (LG)]. Scale=10 �m.


atlantica (Jørgensen 2000a). None of theseepithets match this species. The most obvi-ous difference between all these species andP. effilata is the presence of large oxalatecrystals (crystallocumuli) in their thalli andin the thallus layer covering the perithecia;indeed, all these species belong to Porina s.str. sensu Harris (1995: 170, 172–175) or tothe Porina nucula group sensu Lücking(2004: 410–412). However, the perithecia ofP. effilata can reach full maturity (i.e. pro-ducing mature ascospores) while still deeplyimmersed in the thallus, and thus with onlytheir upper part visible; even when sessileand subglobose, their bases can still becovered by a thin thallus layer; these twofeatures can be confusing. Moreover, thelong and tapering ascospores of both P.guaranitica and P. heterospora are quite simi-lar to those of P. effilata and we suspect thatSwinscow (1962: 47–49) had been confusedby such a strong convergence.

In the field, Porina effilata can be confusedwith Belonia lumbrispora Etayo (Etayo 1996)with which it often grows in the CanaryIslands. The perithecia of the latter arehowever slightly smaller, more yellowish ororange and are not pilose. Examinationunder the microscope immediately confirmsthat Belonia has a hamathecium filled withyellow oily droplets and ascospores thatare multiseptate (40–55 septa), filiform,flexuose, very narrow, and measuring125–160�3–4 �m.

The specimens from Macaronesia differsomewhat from those from the British Isles,mainly Ireland. All major characters that arediagnostic for the species are shared by bothpopulations, but there are slight but signifi-cant differences in the size of perithecia andascospores. This matter has not beenstudied in detail but can be demonstrated bythe following comparison: A. M. Brand13302 (hb Brand) from La Palma (CanaryIslands) has perithecia always distinctlypilose, 0·4–0·5 mm diam., and ascospores(60–)62–70·8–81(–85) �m long (n=22)(Fig. 6B), while the collection by P. W.James, coll. 11 viii 1966 (BM) fromGlengarriff, West Cork, Ireland, has smoothor rarely pilose perithecia, 0·5–0·6 mm

diam., and ascospores (76–)80–87·6–94(–104) �m (n=16) (Fig. 6C). Ascosporeshave been measured in standard conditions(mounted in water; only those expelled fromasci under gentle pressure measured). Wewould not be surprised that future studies,based on more detailed analysis of the vari-ation (including molecular techniques),might indicate two different taxa areinvolved.

Ecology and distribution. Porina effilata hasbeen found in Madeira and in the CanaryIslands (Tenerife, Gomera and La Palma),always in rather preserved stands of thelaurisilva where it can grow on bark as wellas on twigs. It is rare but, as it is an incon-spicuous species, it may have been over-looked. It is also known from SW Ireland,Wales, and N Devon in England, where it isalso quite rare and grows on old, mossy barkof Quercus, usually at the base of the trees, inhighly preserved and humid localities. Bothcollections from Wales and one from Irelandare from bryophytes growing over rocks insheltered conditions, and thus represent theonly localities where P. effilata is not corti-colous; one of them is the type locality ofthe recently described Biatora britannicaPrintzen, Lumbsch & Orange (Printzen et al.2001). An additional locality is known inPortugal in the famous locality at themonastery of Sintra where the species isfound in much more artificial conditions(bole of Cupressus in parkland).

Specimens examined. Great Britain: England: V.C. 4,North Devon: Bideford, Clovelly, woodland below(NW of) Gallantry Bower, on mossy base of largeQuercus, 45 m, 1994, B. J. Coppins 16465 & A. M.O’Dare (E). Wales: V.C. 46, Cardiganshire: nearCardigan, Coedmor National Nature Reserve, SE ofCoedmore house, on bryophytes on vertical rock facesheltered by ivy, 1996, A. Orange 10998 (NMW). V.C.48, Merioneth: Talsarnau, Bryn Bwbach, CeunantCoch, on slightly calcareous rock face in woodland,2002, A. Orange 13840 (NMW).—Ireland: V.C. H2,North Kerry: Killarney, Cromaglown, on trees, viii1935, D. A. Jones s. n. (BM); Eagle’s Nest, shady ledgeon shale cliff, on bryophytes, 26 vi 1961, T. D. V.Swinscow s. n. (BM, 2 specimens); near Killarney,Eagle’s Nest Mountain, on Quercus in wood, 14 viii1966, P. W. James s. n. (BM); Torc Mountain, oak inshade on west-facing slope, 31 vii 1965, T. D. V.Swinscow s. n. (BM); Killarney, Muckross, on Quercus,


14 viii 1966, P. W. James s. n. (BM, E); Killarney Lake,Dinish Island, Camillan wood, on oak, 17 ix 1982, P.W. James s. n. (BM), P. M. Jørgensen 9121 (BG—L/48007) and s. n. (E); Killarney, Meeting of the Waters,N of old Weir Bridge, on old sheltered Quercus at shore,2000, A. M. Brand 40242 (hb Brand). V. C. H3, WestCork: Glengarriff, road to Barley Lake, on shaded boleof Quercus, 11 viii 1966, P. W. James s. n. (BM).—Portugal: W of Lisboa, Sintra, near Capuchosmonastery, on old Cupressus, 250 m, 2003, A. M. Brand49782 (hb Brand).—Canary Islands: La Palma:4·5 km WSW of Los Sauces, Los Tilos, laurisilva,narrow cleft with path along N facing rock sheer, onLaurus azorica and Ocotea foetens, 750–800 m, 1999, P.van den Boom 22240 & 22273 (hb van den Boom); ibid.,narrow cleft with path over Bco del Aqua, mixed treesnear bridge, on tree, 800 m, 1999, P. van den Boom22285 (hb van den Boom); ibid., gorges profondes aveclaurisilve, 600–700 m, sur arbre, iv 2004, E. Sérusiaux s.n. (LG, 5 collections); 8.5 km N of Santa Cruz, Bco laGalga, Cubo de la Galga, laurisilva forest in cleft,550 m, 1999, P. van den Boom 22457 (hb van denBoom); ibid., on bark of young trees of Lauraceae inwood, 580 m, 1986, A. M. Brand 13302 (hb Brand);ibid., 500–600 m, on twigs, iv 2004, E. Sérusiaux s. n.(LG). Tenerife: Laurisilve de Monte del Agua, cheminau départ de Erjos, vers Las Portelas, laurisilveen mosaïque avec des fourrés de Erica, sur troncs,900 m, 27 ii et 2 iii 1997, E. Sérusiaux s. n. (LG, 3collections).—Madeira: Casa das Queimadas, cheminvers Caldeirâo Verde, laurisylve�dégradée, 850–900 m, v 1992, E. Sérusiaux s. n. (LG); S de Seixal,Chão da Ribeira, le long du Riba da Seixal, plus hautque l’élevage de truites, laurisilve en bord de torrent, surtronc, 500 m, vii 2003, E. Sérusiaux s. n. (LG, 2collections).

Porina fortunata P. M. McCarthy &Etayo

Lichenologist 34: 199 (2002); type: Canary Islands,Gomera, La Meseta de Vallehermoso, cliff of the‘‘Cueva Encantada’’, on shaded basalt in laurisilva,720 m, 22 vii 2000, J. Etayo 17823 & A. Fernández(TFC—holotypus, not seen; CANB, hb Etayo—isotypi,not seen).

This saxicolous species has been recentlydescribed from shaded basalt rocks in thelaurisilva of Gomera in the Canary Islands(McCarthy & Etayo 2002), and is herereported from similar habitats in La Palma(collection checked by P. M. McCarthy).Porina fortunata has hemispherical to sub-globose, small [(0·22–)0·28(–0·36) mmdiam.], dark reddish brown to �blackperithecia and ascospores 3(–7)-septate,narrowly oblong to almost cylindrical with

rather rounded ends, 18–25–33�2·5–3–4 �m.

Specimen examined. Canary Islands: La Palma:8·5 km N of Santa Cruz, Barranco La Galga, laurisilvaforest in cleft with volcanic rockface, on shaded basaltboulder, 500 m, 1999, P. van den Boom 22447 (LG, hbvan den Boom).

Porina leptospora (Nyl.) A. L. Sm.

Monogr. Brit. Lich. 2: 338 (1911).—Verrucarialeptospora Nyl., Flora 47: 487 (1864).

Porina olivacea (Pers.) A. L. Sm. var. leptospora (Nyl.)Keissler, Rabenhorst Kryptogamen Flora, Band 9, Abt. 1,Teil 2: 318 (1938).

Porina borreri (Trevis.) D. Hawksw. & P. James var.leptospora (Nyl.) D. Hawksw., Lichenologist 24: 367(1992). Type: Ireland: Kerry, Killarney, Dinish, on Ilex,I. Caroll (?BM—holotypus; not seen).

(Fig. 5D)

This taxon is either considered to be syn-onymous with Porina borreri (Trevis.) D.Hawksw. & P. James (Santesson et al. 2004:271, sub Pseudosagedia borreri), or a variety[var. leptospora (Nyl.) D. Hawksw.; Purviset al. 1992: 490]. In corticolous material ofPorina with black perithecia and 7-septateascospores in Macaronesia, we encounteredonly populations with narrow, almost cylin-drical ascospores, rarely tapering towardsone end that measure 37–50�2·5–3·5(–4·0) �m. In material identified as P. borreriin continental Europe, ascospores are fusi-form or clavate-fusiform and measure 22–33(–35)�3–5 �m (Fig. 5E & F). We didnot find any specimens with intermediateascospores and thus believe the species rankis more appropriate for these taxa.

Porina leptospora occurs in the laurisilvain Macaronesia (La Palma and Tenerife,Canary Islands and São Miguel in theAzores) and, to our knowledge, is knownwith certainty only from SW Ireland andDevon in Western Europe. Although wehave not seen the material, we assume thatthe reports of P. borreri from Madeira byKalb & Hafellner (1992: 81–82) and LosTilos in La Palma by Etayo (1996: 156, subPseudosagedia borreri) belong to P. leptospora.

Selected specimens examined. Canary Islands: LaPalma: Los Tilos, W de Las Lomados, laurisilve


dominée par Ocotea foetens, riche en Hedera canariensis,sur tronc, 600–700 m, vii 1997, E. Sérusiaux s. n. (LG);ibid., sur un vieux Ocotea, iv 2004, E. Sérusiaux s. n.(LG). Tenerife: laurisilve de Monte del Agua, chemin audépart de Erjos vers Las Portelas, sur tronc d’Erica, 27ii 1997, E. Sérusiaux s. n. (LG).—Azores: São Miguel:Praia, Caminho do Praia to Lagoa do Fogo, shadyhollow, on Myrica faya, 120 m, 2003, F. Berger 17801(hb Berger); Lagoa Furnas, Uferweg an NW Ecke, aufAlnus cordifolia & Quercus sp., 280 m, 2001 & 2003, F.Berger 15974, 17813, 17814 (hb Berger).

Selected specimens examined for Porina borreri:Belgium: prov. Namur: 1·2 km S of Furfooz, Parcnational de Haute Recène, S-exposed slope, on bigAcer, 160 m, 1993, P. van den Boom 13835 (LG).—France: dépt. Pyrénées-Atlantiques: Ste Engrâce, gorgesde Kakouetta, fourrés de Buxus très humides à l’entréedes gorges, sur Buxus, 10 vii 1989, E. Sérusiaux s. n.[with P. W. James, F. Rose & J. Vivant] (LG).

Porina ocoteae Brand & Sérus. sp.nov.

A speciebus generis in Europa et Macaronesia differta sporis elongatis-fusiformibus, 7–11(–13)-septate,(42–)44–50(–53)�(3·5–)4–5 �m.

Typus: Canary Islands, La Palma, W de Los Sauces,‘‘Los Tilos’’, gorges profondes avec laurisilve etprésence de vieux fûts d’Ocotea, base de tronc d’Ocotea,alt. 600–700 m, April 2004, E. Sérusiaux s. n. (LG—holotypus).

(Figs 1E & F, 6D & E, 7C)

Thallus epiphloeodal, able to overgrowliverworts and mosses growing on the bark,very rarely (one specimen seen) on basalticrock at the base of a tree on which thespecies was abundant, crustose, shiny greento grey-brown, or sometimes pale brownishor olive-grey, usually maculate with blackirregular patches, adherent to the bark butfrequently forming bullate portions, bulgesor ridges, and thus appearing to peel offfrom the bark, locally with oxalate crystalsdeposited in globose or irregular masses(crystallocumuli sensu Hafellner & Kalb1995: 163–164); prothallus almost alwayspresent, to 0·5 mm wide, black or bluishblack, sometimes covered by a thin, whiteand conspicuous hyphal layer that can givethe prothallus a fibrous or cottony appear-ance. Isidia locally present, sometimescommon, especially on thalli without peri-thecia and on apparently stressed specimens,

simple or coralloid, shiny or with an erodedsurface, sometimes globose and constrictedseveral times along their length, brittle andeasily removed, concolorous with the thallusor paler, 0·1–1·0 mm high.

Perithecia abundant or absent, especiallyabsent on thalli with well-developed isidia,hemispherical to subglobose, very rarelyconstricted at the base, 0·5–0·6(–0·7) mmdiam., typically covered by a thallus layercontaining masses of oxalate crystals andthus appearing as included in a large thalluswart; ostiolar region slightly depressed ornot, concolorous with the thallus or paleorange to reddish, sometimes brownish.Involucrellum typically pale orange and K+

F. 7. Cross-sections through perithecia of Porinaspecies. A, P. atlantica [Madeira, Chão da Ribeira, vii2003, E. Sérusiaux s. n. (LG)]; B, Porina effilata,perithecium still covered with a thallus layer on its rightside and with its left side emerged from the thallus andslightly pilose on its upper part [Gomera, El Cedro, 26vii 1994, E. Sérusiaux s. n. (LG—holotypus)]; C, P.ocoteae [La Palma, Los Tilos, iv 2004, E. Sérusiaux s. n.

(LG—holotypus)]. Scale=100 �m.


bright orange to reddish, best developed inthe upper half of the perithecium, c. 25 �mthick. Excipulum pale yellow, c. 10 �m thick.Paraphyses abundant, simple or rarelybranched, especially near the base, notinflated at apices, c. 1·5 �m thick. Ascicylindrical-clavate, 150–170�10–15 �m.Ascospores 8/ascus, elongate-fusiform, some-times slightly tapering towards one end,straight or slightly curved, 7–11(–13)-septate, (42–)44–50(–53)�(3·5–)4–5 �m,without a perispore.

Pycnidia rare, immersed in the thallus,c. 0·2 mm in diam., with a hyaline orpale orange, K+ bright orange wall.Pycnidiospores ellipsoid, hyaline, simple,2·5–3·5�1·0–1·5 �m.

Notes. In Macaronesia, this species iseasily distinguished from all other corti-colous Porina with large perithecia by itsshiny green to grey-brown thallus, usuallymaculate with black irregular patches, peel-ing off from the bark and frequently formingbullate portions, bulges or ridges, a typicalblack or bluish black prothallus, and narrowascospores (not exceeding 5 �m wide).

Porina ocoteae belongs to the P. mastoideagroup (referred to the genus Clathroporina byHarris 1995: 171). Closely related speciesinclude the pantropical P. africana Müll.Arg. and P. tetracerae (Ach.) Müll. Arg.(Fig. 6F & G). The former has peritheciawith a typically black periostiolar regionand less septate ascospores [(5–)7(–9):McCarthy 2001: 113] and the latter also hasperithecia with a dark periostiolar region andregularly 7-septate ascospores 24–44 �mlong (McCarthy 2001: 150). The materialfrom Florida (USA) identified by Harris(1995: 171) as P. tetracerae has longerascospores (35–50 �m). In any case, P.ocoteae consistently has narrower ascosporesand most have more than 7 septa, making itsdistinction from P. africana and P. tetraceaeeasy. Dr P. M. McCarthy has studied severalof our collections and confirmed (in litt., 4.2003) that they represent an undescribedspecies.

Ecology and distribution. The new species isnamed after its principal phorophyte, thelaurel tree Ocotea foetens (Ait.) Benth. &Hook. f. (‘‘el til’’ in Spanish). On the islandof La Palma (Canary Islands) in the twofamous sites of Los Tilos and Cubo de LaGalga, Ocotea foetens forms a few spectaculargroves of old trees, becoming multi-stemmed by self-coppicing and with largeand complex bases, which develop woodrolls expanding downwards and on whichthis new species of Porina develops con-spicuous populations. Porina ocoteae hasnever been observed on other tree species,even at Los Tilos; it is however able to growon basaltic stones embedded in the bases ofOcotea trees on which it is present. Fullydeveloped and fertile collections are knownonly from Los Tilos; mostly sterile speci-mens are known from the other localities inLa Palma and on São Jorge in the Azores.Accompanying species in Los Tilos includePorina aenea, P. leptospora and the very rareStrigula brevis (Roux & Sérusiaux 2004:53–55). Easily detected because of its shinygreen thallus with a conspicuous black pro-thallus, Porina ocoteae must be consideredas a very rare species as it has not beenobserved elsewhere in the Canary Islands,nor in Madeira. Its status in the Azores mustbe further studied.

Specimens examined. Canary Islands: La Palma: LosTilos (W de Las Lomados), laurisilve dominée parOcotea foetens, riche en Hedera canariensis, on bark,600–700 m, vii 1997, E. Sérusiaux s. n. (LG, 2 collec-tions); ibid., 800–850 m, vii 1997, E. Sérusiaux s. n.(LG); ibid. narrow cleft with path along N facing rocksheer, on Ocotea, 700 m, 1999, P. van den Boom 22174(hb van den Boom); 3·5 km WSW of Los Sauces, Nslope of Barranco del Agua, at the base of Ocotea inlaurel wood, 530 m, 1986, A. M. Brand 13599 (hbBrand); 8·5 km N of Santa Cruz, Bco La Galga,laurisilva forest in cleft, on Ocotea, 500 m, 1999, P. vanden Boom 22424 (hb van den Boom); La Galga, gorgesau lieu-dit ‘‘Cubo de La Galga’’, 500–600 m, laurisilvesur flancs pentus, iv 2004, E. Sérusiaux s. n. (LG).—Azores: São Jorge: Weg Ribiera da Cedro – Faja doAlem, Hortensienhecke, auf abgestorbener Hydrangia,450 m, 2001, F. Berger 15771 (hb Berger).

Selected specimens examined for other species mentioned.Porina africana Müll. Arg.: Guatemala: Dept. BajaVerapaz, SSE of Coban, SSE of Purulhá, BiotopeMario Dary Rivera, 15(13.2#N 90(13.9#W, NEexposed slope with tropical rain forest, on trunk of small


tree, 1850 m, 2004, P. van den Boom 33423 (hb van denBoom, LG).

Porina tetracerae (Ach.) Müll. Arg.: Rwanda: prov.Cyangugu: Cyamudongo forest, 02(33#50·6$S,28(58#94·9$E, lower montane forest with very tall trees

of Newtonia buchananii, Entandrophragma excelsum, etc.,on small trunks and lianas, 2000 m, 30 iii 2005, E.Sérusiaux s. n. [Field trip to Rwanda with Damien Ertz,Eberhard Fischer & Dorothee Killmann, 2005] (LG).

Key to the species of Porina in Macaronesia

1 On living leaves; ascospores 3-septate . . . . . . . . . . . . . . . . . . . . . . . 2On bark; ascospores with 3 or more septa . . . . . . . . . . . . . . . . . . . . 4On rocks; ascospores with 3 or more septa . . . . . . . . . . . . . . . . . . . . 9

2(1) Perithecia black, usually lens-shaped, rarely hemispherical . P. oxneri R. Sant.[very rare: reported only once from Gomera, Canary Islands by Etayo (1998: 102), sub Pseudosagediaobsoleta (Oksner) Hafellner & Kalb; not checked by us]

Perithecia never ‘‘pure’’ black, always with a reddish tinge, always constricted at thebase . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

3(2) Perithecia 0·2–0·3 mm diam., subglobose or almost so, brownish red, at least whenyoung partly covered by hyphal tissue; algal cells typically present in a layerbetween the inner and outer walls of the perithecium . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . P. hoehneliana (Jaap) R. Sant.[very rare: known only from two localities in Madeira, i.e. the laurisilva of Riba da Seixal at300–400 m, where it is abundant, and Chão de Louros where it was found in 1951 by C. Tavares butnot seen there since]

Perithecia 0·1–0·2 mm diam., globose or vertically elongated, dark red to darkreddish brown; no hyphal tissue covering the perithecia and no algal cells presentbetween the perithecial walls . . . . . . . . . . . P. leptosperma Müll. Arg.[known only from Madeira where it is abundant in the laurisilva, even when heavily disturbed]

4(1) Thallus bluish grey or grey, rather pinkish when fresh, with abundant, simple tocoralloid isidia which usually have hyaline, apical ‘hairs’; perithecia rare orabsent; ascospores ellipsoid, 9–11-septate, c. 40–65�9–15 �m . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . P. coralloidea P. James[=Zamenhofia coralloidea (P. James) Clauzade & Cl. Roux]

[reported from Tenerife and Gomera in the Canary Islands, and the Azores archipelago; mostlyfound on old trunks of arborescent Erica at the edge of laurisilva; never abundant]

Thallus never bluish grey (in P. ocoteae, a bluish grey prothallus is present, and inP. effilata, the thallus can sometimes have a bluish tinge) and never with isidiaterminated with hyaline ‘hairs’ . . . . . . . . . . . . . . . . . . . . . . . . . 5

5(4) Perithecia black or almost so . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Perithecia never black or almost so, ostiolar region sometimes dark brown or rarely

blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

6(5) Ascospores 3-septate, ellipsoid and rarely exceeding 20 �m in length . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . P. aenea (Wallr.) Zahlbr.

[on trees; probably rare in Macaronesia: only reported from Tenerife by Gil González et al. (1990:106) and La Palma by Etayo (1996: 156); a single collection from La Palma, Los Tilos seen by us.The reports of P. chlorotica on trees in Tenerife by Champion (1976: 30) and Gomera by Etayo(1998: 102) may refer to this species]

Ascospores 7-septate, almost cylindrical and always exceeding 20 �m in length .. . . . . . . . . . . . . . . . . . . . . . . . . P. leptospora (Nyl.) A. L. Sm.

[on trees; specimens seen by us from La Palma and Tenerife in the Canary Islands and São Miguelin the Azores; reports of P. borreri from Madeira (Kalb & Hafellner 1992: 81–82) and La Palma(Etayo 1996: 156) not checked but assumed to represent this species]


7(5) Perithecia subglobose, 0·3–0·45(–0·5) mm diam., pale orange to orange, some-times pinkish, sometimes distinctly and finely greyish whitish tomentose in theupper half, rarely slightly verruculose, but never containing agglomerations ofoxalate crystals; ascospores narrowly fusiform, slightly but distinctly inflated inthe upper half and tapering towards the proximal end, straight or most usuallyslightly curved, (7–)9–11(–13)-septate, (65–)68–83(–85)�7–11 �m, with a thinbut distinct perispore . . . . . . . . . . . . . . . . P. effilata Brand & Sérus.[on bark and twigs in the laurisilva of Madeira and Gomera, La Palma and Tenerife in the CanaryIslands; rare but possibly overlooked]

Perithecia hemispherical to subglobose, containing agglomerations of oxalatecrystals in their outer walls, always larger when mature . . . . . . . . . . . . 8

8(7) Thallus rugose to verrucose, sometimes wrinkled, sometimes with isidioid out-growths, without a prothallus; perithecia 0·5–0·9(–1·1) mm diam.; ascosporesfusiform, 7–9(–13)-septate, 44–80�7–13 �m, plus a distinct, 2–3 �m thickperispore . . . . . . . . . . . . . . . . . . P. atlantica (Erichsen) P. M. Jørg.[on bark in the laurisilva of Madeira (locally very abundant) and Gomera, La Palma and Tenerife inthe Canary Islands]

Thallus usually maculate with black irregular patches, adherent to the bark butpeeling off and forming bullate portions, bulges or ridges, locally isidioidoutgrowths present, sometimes common, especially on thalli without perithecia,with a black or bluish black prothallus; perithecia 0·5–0·6(–0·7) mm diam.;ascospores elongate-fusiform, 7–11(–13)-septate, (42–)44–50(–53)�(3·5–)4–5 �m, without a perispore . . . . . . . . . . . . . P. ocoteae Brand & Sérus.[known only at the bases of old boles of Ocotea foetens, in localities with well-preserved laurisilva inLa Palma, Canary Islands—the most important being the most famous laurisilva of ‘‘Los Tilos’’, andfrom São Jorge in the Azores—Endemic]

9(1) Perithecia either yellow-orange or covered by a thallus layer but with the ostioleusually pale orange, pinkish to brownish; ascospores at least 7-septate . . . 10

Perithecia black or dark reddish brown; ascospores 3 or 7-septate . . . . . . 1110(9) Perithecia 0·3–0·4 mm diam., yellow-orange to brownish red, not covered by a

thallus layer containing oxalate crystals; thallus thin and smooth; ascospores7-septate, 42–52�12–15 �m, plus a c. 2 �m thick perispore . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . P. ahlesiana (Körb.) Zahlbr.[known from shaded basaltic boulders in laurisilva; reported from Gomera, Canary Islands byMcCarthy & Etayo (2002) and Graciosa in the Azores by Aptroot & Rodrigues (2005), from LaPalma and São Jorge in the Azores (LG, hb Berger); specimens from Gomera and Graciosa not seenby us]

Perithecia 0·5–0·9(–1·1) mm diam., usually distinctly constricted at their base,typically covered by a thallus layer containing agglomerations of oxalate crystalsand thus concolorous with the thallus; ostiole usually pale orange, pinkish tobrownish; thallus rather thick and rugulose-verruculose; ascospores fusiform,7–9(–13)-septate, 44–80�7–13 �m, plus a distinct, 2–3 �m thick perispore .

. . . . . . . . . . . . . . . . . . . . . . . P. atlantica (Erichsen) P. M. Jørg.[normally a corticolous species but found once on basaltic boulders inside laurisilva at La Palma,Canary Islands]

11(9) Perithecia hemispherical to subglobose, small [(0·22–0·28(–0·36) mm diam.], darkreddish brown to �black; ascospores 3(–7)-septate, narrowly oblong to almostcylindrical with rather rounded ends, 18–33�2·5–4 �m . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . P. fortunata P. M. McCarthy & Etayo[known from shaded basaltic boulders in laurisilva; only two localities, in Gomera and La Palma,Canary Islands—Endemic]

Perithecia usually hemispherical and rather immersed in the thallus, rarelysubglobose, never dark reddish brown, always black . . . . . . . . . . . . 12


12(11) Perithecia 0·2–0·3 mm diam., very rarely aggregated; ascospores 3-septate, 16–25(–32)�4–6 �m . . . . . . . . . . . . . . P. chlorotica (Ach.) Müll. Arg.[reported from the Azores (Degelius 1941: 8), Madeira (Tavares 1952: 319), Gomera (McCarthy &Etayo 2002), and from La Palma and Tenerife, at: www.gobcan.es/medioambiente/biodiversidad/ceplam/bancodatos/libro.html]

Perithecia 0·4–0·6 mm diam., sometimes aggregated into clusters of 2–4; as-cospores 7-septate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13(12) Ascospores fusiform to oblong-cylindrical with rounded ends, (26–)30–45�5–6 �m; thick (up to 5 �m) perispore present but not easily seen, usually associatedwith immature ascospores only . . . . . . . . . P. guentheri (Flot.) Zahlbr.[known from shaded basaltic boulders, usually in laurisilva, in Gomera and La Palma, CanaryIslands; only specimens from La Palma seen by us—report from La Gomera at: www.gobcan.es/medioambiente/biodiversidad/ceplam/bancodatos/libro.html; recently reported from Graciosa in theAzores by Aptroot & Rodrigues (2005)]

Ascospores narrow-fusiform to cylindrical, with rounded ends but usually slightlytapering towards one, 42–55�3–4 �m; perispore not seen . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .P. curnowii A. L. Sm.[reported here from coastal localities, at low elevations in São Jorge and Flores, Azores; coll. in hbBerger]

Excluded species

Porina glabra (A. Massal.) Zahlbr.: thisspecies is included in the checklist oflichens and lichenicolous fungi in theCanary Islands, available at the addressbelow. The name refers to Strigula glabra(A. Massal.) V. Wirth, a species forwhich no material has been seen fromMacaronesia (Roux & Sérusiaux 2004).www.gobcan.es/medioambiente/biodiversidad/

ceplam/bancodatos/libro.htmlPorina rosei Sérus.: the doubtful report of a

sterile collection of that species (Sérusiaux1991) has been assumed to be actuallyone of Porina isidiata Kalb & Hafellner(Kalb & Hafellner 1992). We here con-firm that the specimen (LG!) is close tobut not unequivocally conspecific with P.rosei and is not an isidiate form of Porinaatlantica.

P. semecarpi Vain.: the only collection of thatspecies mentioned from Madeira belongsto P. hoehneliana (Sérusiaux 1996: 225);the world distribution map of the speciespublished by McCarthy (2003: 89) shouldthus be corrected.

We thank the curators of the following herbaria for theloan of material in their care: BG, BM, E, G, NMW, Sand hb Kalb. Dr P. Diederich kindly made his materialof Porina from Papua New Guinea available to us andDr Robert Lücking kindly provided relevant data on the

type collection of Porina nucula Ach. in H and NY. DrP. M. McCarthy examined several specimens of oursfrom Macaronesia and made available his valuablenotes and observations: we owe him a great debt; hefurther reviewed carefully the ms as a referee andprovided additional useful comments. We also thankAlan Orange, the second referee, for his interestingcomments and suggestions.

R

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Accepted for publication 25 August 2006


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