10
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Two New Species of Caloplaca (Teloschistaceae) from the Southern Hemisphere Author(s): Ingvar Kärnefelt, Sergey Kondratyuk, Ulrik Søchting, Patrik Frödén, and Ulf Arup Source: The Bryologist, 105(3):301-309. 2002. Published By: The American Bryological and Lichenological Society, Inc. DOI: http://dx.doi.org/10.1639/0007-2745(2002)105[0301:TNSOCT]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1639/0007-2745%282002%29105%5B0301%3ATNSOCT %5D2.0.CO%3B2 BioOne (www.bioone.org ) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use . Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.

Two New Species of Caloplaca (Teloschistaceae) from the Southern Hemisphere

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors nonprofit publishers academic institutions researchlibraries and research funders in the common goal of maximizing access to critical research

Two New Species of Caloplaca (Teloschistaceae) from the Southern HemisphereAuthor(s) Ingvar Kaumlrnefelt Sergey Kondratyuk Ulrik Soslashchting Patrik Froumldeacuten and Ulf ArupSource The Bryologist 105(3)301-309 2002Published By The American Bryological and Lichenological Society IncDOI httpdxdoiorg1016390007-2745(2002)105[0301TNSOCT]20CO2URL httpwwwbiooneorgdoifull1016390007-2745282002291055B03013ATNSOCT5D20CO3B2

BioOne (wwwbiooneorg) is a nonprofit online aggregation of core research in the biological ecological andenvironmental sciences BioOne provides a sustainable online platform for over 170 journals and books publishedby nonprofit societies associations museums institutions and presses

Your use of this PDF the BioOne Web site and all posted and associated content indicates your acceptance ofBioOnersquos Terms of Use available at wwwbiooneorgpageterms_of_use

Usage of BioOne content is strictly limited to personal educational and non-commercial use Commercial inquiriesor rights and permissions requests should be directed to the individual publisher as copyright holder

0007-274502301ndash309$1050

THE BRYOLOGISTA JOURNAL OF BRYOLOGY AND LICHENOLOGY

VOLUME 105 NUMBER 3FALL 2002

The Bryologist 105(3) pp 301 309Copyright q 2002 by the American Bryological and Lichenological Society Inc

Two New Species of Caloplaca (Teloschistaceae) from the Southern Hemisphere

INGVAR KARNEFELT

The Botanical Museum University of Lund Ostra Vallgatan 18 S-223 61 Lund Sweden e-mailIngvarKarnefeltbotmusluse

SERGEY KONDRATYUK

N G Kholodny Institute of Botany Tereshchenkivsrsquoka 2 252601 Kiev-4 Ukraine e-mail skondrbotankievua

ULRIK SoslashCHTING

Department of Mycology Botanical Institute Oslash Farimagsgade 2D DK-1353 Copenhagen Denmark e-mailulriksbotkudk

PATRIK FRODEN AND ULF ARUP

The Botanical Museum University of Lund Ostra Vallgatan 18 S-223 61 Lund Sweden

Abstract Caloplaca gyalectoides S Kondratyuk amp Karnefelt from the Cape Province SouthAfrica and Caloplaca chilensis S Kondratyuk Karnefelt Froden amp Arup with an associatedparasymbiont Arthonia tetraspora S Kondratyuk amp Karnefelt occurring mainly along the coast incentral Chile are described as new Possible associated taxa are briefly discussed Distributionmaps are provided for all three species

The genus Caloplaca Th Fr comprises a largenumber of species presumably over 800 occurringin many different biota and habitats over the worldSeveral more or less distinctive groups of speciescan be recognized although intermediate forms oc-cur in many cases (Clauzade amp Roux 1985 Kar-nefelt 1989) There is no critical checklist of theentire genus however a list of names has been pro-duced by Cliff Wetmore (httpwwwtcaumneduwetmoreAllCaltophtm) The entire genus hasnever been monographed but smaller groups ofspecies from different geographical regions havebeen revised eg Eurasia and North America (Arup1995 Hansen et al 1987 Navarro-Rosines amp Hla-dun 1996 Poelt amp Hinteregger 1993 Wetmore2001 Wetmore amp Karnefelt 1998 2000) The moretroublesome discussion related to the genus other-wise concerns the distinction between Caloplacaand Xanthoria where we never seem to have ar-rived at more than a temporary solution (Karnefelt1989)

During the work with xanthorioid lichens fromthe southern hemisphere (Karnefelt et al 2002) andfieldwork by Froden and Arup in South Americatwo undescribed species belonging to the genusCaloplaca were found They are described here andnamed C gyalectoides and C chilensis Further-more a lichenicolous fungus from the Arthoniadestruens aggregate appeared on the thallus andthalline margins of C chilensis

MATERIAL AND METHODS

The study presented here is based mainly on herbariummaterial kept in B BCRU BG BM C GZU H KOELN KWLD MIN S SGO TNS and UPS For comparison materialfrom many other herbaria was also consulted eg CBGCOLO herb Kalb LE LEM MEL and MVM

Anatomy Fragments of lichens were sectioned with aKryomat Leitz freezing microtome and sections weremounted in water or in lactophenol cottonblue Anatomi-cal structure and hymenial characters were studied with aZeiss Axioscope light microscope photomicrographswere made with an Olympus DP 11 digital camera

302 [VOL 105THE BRYOLOGIST

Secondary metabolites Extractions of secondary me-tabolites and HPLC analyses were made according toSoslashchting (1997 2001) Quantification of the metaboliteswas based on absorption at 270 nm The proportions ofmetabolites were calculated in relation to the total of allsignificant absorption peaks recorded Thalli and apothe-cia were analyzed separately

THE NEW SPECIES

CALOPLACA GYALECTOIDES S Kondratyuk amp Kar-nefelt sp nov FIGS 1 3

Apothecia primo clausa marginem thallinum roseumtantum monstrantia iis speciei generis Gyalectae sat sim-ilia

TYPE SOUTH AFRICA CAPE PROVINCE Namaqua-land Hondeklip Bay on dead shrubs in the dunes nearthe sea 1953 Ove Almborn 4679 (LD holotype GZU KWPRE UPS isotypes)

Thallus distinctly warted 2ndash5 mm diam 02ndash03mm thick composed of aggregated lobules andgroups of clustered algae or soredia-like structuresgreenish gray Central parts and tips of warts withdistinct thick white pruina (Figs 1ab) Apotheciazeorine 05ndash20(ndash27) mm diam margin distinctlywarted disk whitish-yellowish with thin pruinathalline margin 02ndash03(ndash04) mm thick true exci-ple well-developed ca 90ndash100 mm thick in lateralportion thinner in basal portion composed of truescleroplectenchymatous layer with large white ne-crotic zone formed by bundles of hyphae projectingfrom true exciple thalline exciple composed ofscleroplectenchymatous tissue ca 12ndash18(ndash35) mmthick partly covered by groups of parallel hyphaeforming bundles that become white after dying hy-pothecium ca 15 mm thick hymenium ca 60ndash70mm high asci cylindric ca 55 3 12 mm ascosporesnarrowly ellipsoid 115ndash120(ndash130) 3 (60ndash)65ndash70 mm septum 25ndash30 mm thick All yellow partsK1 red C2 I2

Chemistry The only compound recorded fromthe scant thallus material analyzed with HPLC wasparietin This indicates that no other anthraquinonesare present in significant amounts but it is verylikely that extraction of more material would havedisclosed small amounts of teloschistin fallacinalparietinic acid and emodin thus assigning che-mosyndrome A to Caloplaca gyalectoides(Soslashchting 1997)

Etymology The species epithet refers to thestriking resemblance with species of the genus Gy-alecta

Ecology and distribution Caloplaca gyalecto-ides is so far only known from the type locality inthe Cape Province Namaqualand Hondeklip Baywhere it grew on dead shrubs in the dunes near thesea (Fig 2) It occurred together with Xanthoriaturbinata Xanthodactylon flammeum C cf pyra-

cea C cf herbidella Lecanora sp and Micareasp

Taxonomic notes The apothecia at first devel-op without the characteristic white outgrowths Lat-er these outgrowths are formed by bundles of thick-walled parallel-arranged hyphae (Fig 3) The disccan appear as closed showing only the distinct rose-colored thalline margin quite similar to the Gy-alecta type of apothecia

Caloplaca gyalectoides is an unique species inthe genus due to its gyalectoid pinkish apotheciaOtherwise it reminds one somewhat of C pyraceaa species that is not known from the region (Feurer2001) Dodge (1971) did not discuss a species ofthis appearance from Africa nor does it appear inany modern European flora (Clauzade amp Roux1985 Purvis et al 1992) or in the New Zealandlichen flora (Galloway 1985)

CALOPLACA CHILENSIS Karnefelt S KondratyukFroden amp Arup sp nov FIGS 4ndash6

Caloplacae brattiae similis sed differt sporis majoribusapothecii stipitatis et habitatione corticola

TYPE CHILE Reg IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of routa 5 Dry opencoastal hill with shrubs and boulders On different shrubsand more rarely on cacti Alt 200ndash300 m 2984529 S71819829 W 2001-01-26 Froden amp Arup 1 (LD holotypeC BCRU BG GZU KOELN KW MIN SGO isotypes)

Thallus small 8ndash15(ndash25) mm diam formingwell developed rosettes center covered by apothe-cia or partly verruculose from apothecium initials(Fig 4) lobes well-developed radiating rather nar-row and convex irregularly branched closely ad-pressed to substrate or separating from substrate atlobe tips (05ndash)10ndash30(ndash38) mm long 02ndash04(ndash06) mm wide at unbranched parts yellowish orreddish to brownish orange usually paler yellowto yellowish orange at terminal portions of lobessurface often shiny lobes 150ndash200(ndash250) mmthick attached to substrate by medullary hyphaethat sometimes form rhizine-like structures of con-glutinated hyphae (Fig 5) upper cortical layercomposed of thick-walled irregularly arranged hy-phae 5ndash15 mm and with 15ndash20 mm thick luminaalgal layer composed of clustered trebouxoid algaemedulla rather compact lower cortex absent orpresent where lobes lift from substrate (Fig 6) pro-thallus occasionally present orange thin visiblebetween lobes and at thallus margin pseudocy-phellae sometimes present Apothecia mainly pre-sent in central part often abundant occasionallylacking 04ndash27 mm diam stipitate stipe 03ndash10mm disk concave to plane often flexuose orangeto reddish or brownish orange margin yellowishorange true exciple 50ndash60 mm thick in lateral por-

2002] 303KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 1 Habit of Caloplaca gyalectoides holotype (LD) Scale bars a and b 5 1 mm

tion composed of thick-walled irregular arrangedhyphae thalline exciple very thick 75ndash125 mmyellow to orange mostly paler than and raisedabove or level with disk sometimes with radiatingfurrows hymenium 80ndash100 mm high with yellow-brownish epithecium 5ndash12(ndash20) mm thick hypoth-ecium hyaline 25ndash80 mm thick paraphyses septatesimple or apically branched ca 15 mm thick in the

middle part apically slightly swollen to 3 mmdiam asci cylindric 50ndash80 3 10ndash12 mm asco-spores narrowly ellipsoid (12ndash)13ndash17(ndash19) 3(40ndash)45ndash60(ndash75) mm septum (30ndash)40ndash60(ndash75) mm thick sometimes curved spermogoniaprotruding reddish orange ca 01ndash03 mm widespermatia narrowly ellipsoid-bacilliform 33ndash50 309ndash12 mm Thallus and apothecia K1 red C2 I2

304 [VOL 105THE BRYOLOGIST

FIGURE 2 Known distribution of Caloplaca gyalec-toides

FIGURE 3 Bundles of hyphae on the thalline marginof Caloplaca gyalectoides Scale bar 5 10 mm

Chemistry Caloplaca chilensis contains theanthraquinones parietin and its oxidation productsteloschistin fallacinal parietinic acid and emodinParietin is the dominant compound in the apothecia(89) In the thallus parietin accounts for 55 ofthe anthraquinones and teloschistin fallacinal par-ietinic acid and emodin accounts for 15 264 and 1 respectively This anthraquinone che-mosyndrome agrees with A3 of Soslashchting (1997) Itmust be borne in mind that the calculated propor-tions are not absolute quantities but are based onabsorption at 270 nm

Ecology and distribution Caloplaca chilensisgrows on branches of bushes of various sizes aswell as on small trees and cacti The phorophytespecies does not seem to be important as it wasfound growing on at least six different phorophyteshowever always sun exposed The species occurson both dead and living twigs and branches It ismostly found in localities with dry open shrub veg-etation that is often grazed by sheep or other ani-mals (Fig 7) Even though boulders occur in mostknown localities of C chilensis and these oftenhold a large number of other Caloplaca species Cchilensis was never found on rocks The known lo-calities span from about 25 to 1100 m above sealevel However it is likely that C chilensis occursat even higher altitudes In most known localitiesC chilensis was abundant

A number of other species are often found to-gether with C chilensis eg Candelaria sp Chry-sothrix pavonii (Fr) J R Laundon C cf cerinaTeloschistes chrysophthalmus (L) Th Fr Ramal-ina sp Xanthoria ascendens S Kondratyuck andUsnea spp

Caloplaca chilensis apparently is restricted to thepart of Chile (Fig 8) that is characterized by heavyfogs caused by upwelling cold currents along the

coast The currents outside the coast of the Atacamaproduce a stable mild climate with very little pre-cipitation The lichens there receive their water al-most exclusively from the fogs The effect of thefogs on the lichen communities varies dependingon distance from the sea elevation and topogra-phy Normally the fog comes in from the sea dur-ing the night and disappears around noon the nextday (Rundel 1978) Most of the known localitiesfor the species are hills or mountains near the coastbut two localities differ The first is located in thebottom of an inland valley where the fog movesinland The second one is located on a mountain atthe bottom of this valley about 40 km from thecoast There are however no mountains farther tothe coast blocking the way for the fog which canmove inland in such valleys

The described conditions are typical for an areaalong the coast in Peru and the northern part ofChile It is therefore likely that C chilensis will befound to have a larger distributional area than re-ported here However it was not found around Ar-ica in northern Chile or in southern Peru possiblybecause the climate there is too dry Caloplaca chi-lensis does not match any species on the list ofspecies recorded from Chile (Galloway amp Quilhot2001) or any of the species recorded from the re-gion (Follmann 1962 1967 Follmann amp Redon1972 Rasanen 1936 Zahlbruckner 1925)

Taxonomic notes It is remarkable that such acommon and conspicuous species as C chilensisdoes not seem to have been described earlier or atleast reported under some other name as many li-chenologists have visited some of the known lo-calities eg P N Fray Jorge One possible reasoncould be that it has expanded its distribution rangeonly recently in connection with the drastic and insome cases catastrophic changes in the lichen floraof the coastal mountains that has occurred duringthe last decades (Follmann 1995)

Caloplaca chilensis is morphologically similar to

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

0007-274502301ndash309$1050

THE BRYOLOGISTA JOURNAL OF BRYOLOGY AND LICHENOLOGY

VOLUME 105 NUMBER 3FALL 2002

The Bryologist 105(3) pp 301 309Copyright q 2002 by the American Bryological and Lichenological Society Inc

Two New Species of Caloplaca (Teloschistaceae) from the Southern Hemisphere

INGVAR KARNEFELT

The Botanical Museum University of Lund Ostra Vallgatan 18 S-223 61 Lund Sweden e-mailIngvarKarnefeltbotmusluse

SERGEY KONDRATYUK

N G Kholodny Institute of Botany Tereshchenkivsrsquoka 2 252601 Kiev-4 Ukraine e-mail skondrbotankievua

ULRIK SoslashCHTING

Department of Mycology Botanical Institute Oslash Farimagsgade 2D DK-1353 Copenhagen Denmark e-mailulriksbotkudk

PATRIK FRODEN AND ULF ARUP

The Botanical Museum University of Lund Ostra Vallgatan 18 S-223 61 Lund Sweden

Abstract Caloplaca gyalectoides S Kondratyuk amp Karnefelt from the Cape Province SouthAfrica and Caloplaca chilensis S Kondratyuk Karnefelt Froden amp Arup with an associatedparasymbiont Arthonia tetraspora S Kondratyuk amp Karnefelt occurring mainly along the coast incentral Chile are described as new Possible associated taxa are briefly discussed Distributionmaps are provided for all three species

The genus Caloplaca Th Fr comprises a largenumber of species presumably over 800 occurringin many different biota and habitats over the worldSeveral more or less distinctive groups of speciescan be recognized although intermediate forms oc-cur in many cases (Clauzade amp Roux 1985 Kar-nefelt 1989) There is no critical checklist of theentire genus however a list of names has been pro-duced by Cliff Wetmore (httpwwwtcaumneduwetmoreAllCaltophtm) The entire genus hasnever been monographed but smaller groups ofspecies from different geographical regions havebeen revised eg Eurasia and North America (Arup1995 Hansen et al 1987 Navarro-Rosines amp Hla-dun 1996 Poelt amp Hinteregger 1993 Wetmore2001 Wetmore amp Karnefelt 1998 2000) The moretroublesome discussion related to the genus other-wise concerns the distinction between Caloplacaand Xanthoria where we never seem to have ar-rived at more than a temporary solution (Karnefelt1989)

During the work with xanthorioid lichens fromthe southern hemisphere (Karnefelt et al 2002) andfieldwork by Froden and Arup in South Americatwo undescribed species belonging to the genusCaloplaca were found They are described here andnamed C gyalectoides and C chilensis Further-more a lichenicolous fungus from the Arthoniadestruens aggregate appeared on the thallus andthalline margins of C chilensis

MATERIAL AND METHODS

The study presented here is based mainly on herbariummaterial kept in B BCRU BG BM C GZU H KOELN KWLD MIN S SGO TNS and UPS For comparison materialfrom many other herbaria was also consulted eg CBGCOLO herb Kalb LE LEM MEL and MVM

Anatomy Fragments of lichens were sectioned with aKryomat Leitz freezing microtome and sections weremounted in water or in lactophenol cottonblue Anatomi-cal structure and hymenial characters were studied with aZeiss Axioscope light microscope photomicrographswere made with an Olympus DP 11 digital camera

302 [VOL 105THE BRYOLOGIST

Secondary metabolites Extractions of secondary me-tabolites and HPLC analyses were made according toSoslashchting (1997 2001) Quantification of the metaboliteswas based on absorption at 270 nm The proportions ofmetabolites were calculated in relation to the total of allsignificant absorption peaks recorded Thalli and apothe-cia were analyzed separately

THE NEW SPECIES

CALOPLACA GYALECTOIDES S Kondratyuk amp Kar-nefelt sp nov FIGS 1 3

Apothecia primo clausa marginem thallinum roseumtantum monstrantia iis speciei generis Gyalectae sat sim-ilia

TYPE SOUTH AFRICA CAPE PROVINCE Namaqua-land Hondeklip Bay on dead shrubs in the dunes nearthe sea 1953 Ove Almborn 4679 (LD holotype GZU KWPRE UPS isotypes)

Thallus distinctly warted 2ndash5 mm diam 02ndash03mm thick composed of aggregated lobules andgroups of clustered algae or soredia-like structuresgreenish gray Central parts and tips of warts withdistinct thick white pruina (Figs 1ab) Apotheciazeorine 05ndash20(ndash27) mm diam margin distinctlywarted disk whitish-yellowish with thin pruinathalline margin 02ndash03(ndash04) mm thick true exci-ple well-developed ca 90ndash100 mm thick in lateralportion thinner in basal portion composed of truescleroplectenchymatous layer with large white ne-crotic zone formed by bundles of hyphae projectingfrom true exciple thalline exciple composed ofscleroplectenchymatous tissue ca 12ndash18(ndash35) mmthick partly covered by groups of parallel hyphaeforming bundles that become white after dying hy-pothecium ca 15 mm thick hymenium ca 60ndash70mm high asci cylindric ca 55 3 12 mm ascosporesnarrowly ellipsoid 115ndash120(ndash130) 3 (60ndash)65ndash70 mm septum 25ndash30 mm thick All yellow partsK1 red C2 I2

Chemistry The only compound recorded fromthe scant thallus material analyzed with HPLC wasparietin This indicates that no other anthraquinonesare present in significant amounts but it is verylikely that extraction of more material would havedisclosed small amounts of teloschistin fallacinalparietinic acid and emodin thus assigning che-mosyndrome A to Caloplaca gyalectoides(Soslashchting 1997)

Etymology The species epithet refers to thestriking resemblance with species of the genus Gy-alecta

Ecology and distribution Caloplaca gyalecto-ides is so far only known from the type locality inthe Cape Province Namaqualand Hondeklip Baywhere it grew on dead shrubs in the dunes near thesea (Fig 2) It occurred together with Xanthoriaturbinata Xanthodactylon flammeum C cf pyra-

cea C cf herbidella Lecanora sp and Micareasp

Taxonomic notes The apothecia at first devel-op without the characteristic white outgrowths Lat-er these outgrowths are formed by bundles of thick-walled parallel-arranged hyphae (Fig 3) The disccan appear as closed showing only the distinct rose-colored thalline margin quite similar to the Gy-alecta type of apothecia

Caloplaca gyalectoides is an unique species inthe genus due to its gyalectoid pinkish apotheciaOtherwise it reminds one somewhat of C pyraceaa species that is not known from the region (Feurer2001) Dodge (1971) did not discuss a species ofthis appearance from Africa nor does it appear inany modern European flora (Clauzade amp Roux1985 Purvis et al 1992) or in the New Zealandlichen flora (Galloway 1985)

CALOPLACA CHILENSIS Karnefelt S KondratyukFroden amp Arup sp nov FIGS 4ndash6

Caloplacae brattiae similis sed differt sporis majoribusapothecii stipitatis et habitatione corticola

TYPE CHILE Reg IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of routa 5 Dry opencoastal hill with shrubs and boulders On different shrubsand more rarely on cacti Alt 200ndash300 m 2984529 S71819829 W 2001-01-26 Froden amp Arup 1 (LD holotypeC BCRU BG GZU KOELN KW MIN SGO isotypes)

Thallus small 8ndash15(ndash25) mm diam formingwell developed rosettes center covered by apothe-cia or partly verruculose from apothecium initials(Fig 4) lobes well-developed radiating rather nar-row and convex irregularly branched closely ad-pressed to substrate or separating from substrate atlobe tips (05ndash)10ndash30(ndash38) mm long 02ndash04(ndash06) mm wide at unbranched parts yellowish orreddish to brownish orange usually paler yellowto yellowish orange at terminal portions of lobessurface often shiny lobes 150ndash200(ndash250) mmthick attached to substrate by medullary hyphaethat sometimes form rhizine-like structures of con-glutinated hyphae (Fig 5) upper cortical layercomposed of thick-walled irregularly arranged hy-phae 5ndash15 mm and with 15ndash20 mm thick luminaalgal layer composed of clustered trebouxoid algaemedulla rather compact lower cortex absent orpresent where lobes lift from substrate (Fig 6) pro-thallus occasionally present orange thin visiblebetween lobes and at thallus margin pseudocy-phellae sometimes present Apothecia mainly pre-sent in central part often abundant occasionallylacking 04ndash27 mm diam stipitate stipe 03ndash10mm disk concave to plane often flexuose orangeto reddish or brownish orange margin yellowishorange true exciple 50ndash60 mm thick in lateral por-

2002] 303KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 1 Habit of Caloplaca gyalectoides holotype (LD) Scale bars a and b 5 1 mm

tion composed of thick-walled irregular arrangedhyphae thalline exciple very thick 75ndash125 mmyellow to orange mostly paler than and raisedabove or level with disk sometimes with radiatingfurrows hymenium 80ndash100 mm high with yellow-brownish epithecium 5ndash12(ndash20) mm thick hypoth-ecium hyaline 25ndash80 mm thick paraphyses septatesimple or apically branched ca 15 mm thick in the

middle part apically slightly swollen to 3 mmdiam asci cylindric 50ndash80 3 10ndash12 mm asco-spores narrowly ellipsoid (12ndash)13ndash17(ndash19) 3(40ndash)45ndash60(ndash75) mm septum (30ndash)40ndash60(ndash75) mm thick sometimes curved spermogoniaprotruding reddish orange ca 01ndash03 mm widespermatia narrowly ellipsoid-bacilliform 33ndash50 309ndash12 mm Thallus and apothecia K1 red C2 I2

304 [VOL 105THE BRYOLOGIST

FIGURE 2 Known distribution of Caloplaca gyalec-toides

FIGURE 3 Bundles of hyphae on the thalline marginof Caloplaca gyalectoides Scale bar 5 10 mm

Chemistry Caloplaca chilensis contains theanthraquinones parietin and its oxidation productsteloschistin fallacinal parietinic acid and emodinParietin is the dominant compound in the apothecia(89) In the thallus parietin accounts for 55 ofthe anthraquinones and teloschistin fallacinal par-ietinic acid and emodin accounts for 15 264 and 1 respectively This anthraquinone che-mosyndrome agrees with A3 of Soslashchting (1997) Itmust be borne in mind that the calculated propor-tions are not absolute quantities but are based onabsorption at 270 nm

Ecology and distribution Caloplaca chilensisgrows on branches of bushes of various sizes aswell as on small trees and cacti The phorophytespecies does not seem to be important as it wasfound growing on at least six different phorophyteshowever always sun exposed The species occurson both dead and living twigs and branches It ismostly found in localities with dry open shrub veg-etation that is often grazed by sheep or other ani-mals (Fig 7) Even though boulders occur in mostknown localities of C chilensis and these oftenhold a large number of other Caloplaca species Cchilensis was never found on rocks The known lo-calities span from about 25 to 1100 m above sealevel However it is likely that C chilensis occursat even higher altitudes In most known localitiesC chilensis was abundant

A number of other species are often found to-gether with C chilensis eg Candelaria sp Chry-sothrix pavonii (Fr) J R Laundon C cf cerinaTeloschistes chrysophthalmus (L) Th Fr Ramal-ina sp Xanthoria ascendens S Kondratyuck andUsnea spp

Caloplaca chilensis apparently is restricted to thepart of Chile (Fig 8) that is characterized by heavyfogs caused by upwelling cold currents along the

coast The currents outside the coast of the Atacamaproduce a stable mild climate with very little pre-cipitation The lichens there receive their water al-most exclusively from the fogs The effect of thefogs on the lichen communities varies dependingon distance from the sea elevation and topogra-phy Normally the fog comes in from the sea dur-ing the night and disappears around noon the nextday (Rundel 1978) Most of the known localitiesfor the species are hills or mountains near the coastbut two localities differ The first is located in thebottom of an inland valley where the fog movesinland The second one is located on a mountain atthe bottom of this valley about 40 km from thecoast There are however no mountains farther tothe coast blocking the way for the fog which canmove inland in such valleys

The described conditions are typical for an areaalong the coast in Peru and the northern part ofChile It is therefore likely that C chilensis will befound to have a larger distributional area than re-ported here However it was not found around Ar-ica in northern Chile or in southern Peru possiblybecause the climate there is too dry Caloplaca chi-lensis does not match any species on the list ofspecies recorded from Chile (Galloway amp Quilhot2001) or any of the species recorded from the re-gion (Follmann 1962 1967 Follmann amp Redon1972 Rasanen 1936 Zahlbruckner 1925)

Taxonomic notes It is remarkable that such acommon and conspicuous species as C chilensisdoes not seem to have been described earlier or atleast reported under some other name as many li-chenologists have visited some of the known lo-calities eg P N Fray Jorge One possible reasoncould be that it has expanded its distribution rangeonly recently in connection with the drastic and insome cases catastrophic changes in the lichen floraof the coastal mountains that has occurred duringthe last decades (Follmann 1995)

Caloplaca chilensis is morphologically similar to

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

302 [VOL 105THE BRYOLOGIST

Secondary metabolites Extractions of secondary me-tabolites and HPLC analyses were made according toSoslashchting (1997 2001) Quantification of the metaboliteswas based on absorption at 270 nm The proportions ofmetabolites were calculated in relation to the total of allsignificant absorption peaks recorded Thalli and apothe-cia were analyzed separately

THE NEW SPECIES

CALOPLACA GYALECTOIDES S Kondratyuk amp Kar-nefelt sp nov FIGS 1 3

Apothecia primo clausa marginem thallinum roseumtantum monstrantia iis speciei generis Gyalectae sat sim-ilia

TYPE SOUTH AFRICA CAPE PROVINCE Namaqua-land Hondeklip Bay on dead shrubs in the dunes nearthe sea 1953 Ove Almborn 4679 (LD holotype GZU KWPRE UPS isotypes)

Thallus distinctly warted 2ndash5 mm diam 02ndash03mm thick composed of aggregated lobules andgroups of clustered algae or soredia-like structuresgreenish gray Central parts and tips of warts withdistinct thick white pruina (Figs 1ab) Apotheciazeorine 05ndash20(ndash27) mm diam margin distinctlywarted disk whitish-yellowish with thin pruinathalline margin 02ndash03(ndash04) mm thick true exci-ple well-developed ca 90ndash100 mm thick in lateralportion thinner in basal portion composed of truescleroplectenchymatous layer with large white ne-crotic zone formed by bundles of hyphae projectingfrom true exciple thalline exciple composed ofscleroplectenchymatous tissue ca 12ndash18(ndash35) mmthick partly covered by groups of parallel hyphaeforming bundles that become white after dying hy-pothecium ca 15 mm thick hymenium ca 60ndash70mm high asci cylindric ca 55 3 12 mm ascosporesnarrowly ellipsoid 115ndash120(ndash130) 3 (60ndash)65ndash70 mm septum 25ndash30 mm thick All yellow partsK1 red C2 I2

Chemistry The only compound recorded fromthe scant thallus material analyzed with HPLC wasparietin This indicates that no other anthraquinonesare present in significant amounts but it is verylikely that extraction of more material would havedisclosed small amounts of teloschistin fallacinalparietinic acid and emodin thus assigning che-mosyndrome A to Caloplaca gyalectoides(Soslashchting 1997)

Etymology The species epithet refers to thestriking resemblance with species of the genus Gy-alecta

Ecology and distribution Caloplaca gyalecto-ides is so far only known from the type locality inthe Cape Province Namaqualand Hondeklip Baywhere it grew on dead shrubs in the dunes near thesea (Fig 2) It occurred together with Xanthoriaturbinata Xanthodactylon flammeum C cf pyra-

cea C cf herbidella Lecanora sp and Micareasp

Taxonomic notes The apothecia at first devel-op without the characteristic white outgrowths Lat-er these outgrowths are formed by bundles of thick-walled parallel-arranged hyphae (Fig 3) The disccan appear as closed showing only the distinct rose-colored thalline margin quite similar to the Gy-alecta type of apothecia

Caloplaca gyalectoides is an unique species inthe genus due to its gyalectoid pinkish apotheciaOtherwise it reminds one somewhat of C pyraceaa species that is not known from the region (Feurer2001) Dodge (1971) did not discuss a species ofthis appearance from Africa nor does it appear inany modern European flora (Clauzade amp Roux1985 Purvis et al 1992) or in the New Zealandlichen flora (Galloway 1985)

CALOPLACA CHILENSIS Karnefelt S KondratyukFroden amp Arup sp nov FIGS 4ndash6

Caloplacae brattiae similis sed differt sporis majoribusapothecii stipitatis et habitatione corticola

TYPE CHILE Reg IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of routa 5 Dry opencoastal hill with shrubs and boulders On different shrubsand more rarely on cacti Alt 200ndash300 m 2984529 S71819829 W 2001-01-26 Froden amp Arup 1 (LD holotypeC BCRU BG GZU KOELN KW MIN SGO isotypes)

Thallus small 8ndash15(ndash25) mm diam formingwell developed rosettes center covered by apothe-cia or partly verruculose from apothecium initials(Fig 4) lobes well-developed radiating rather nar-row and convex irregularly branched closely ad-pressed to substrate or separating from substrate atlobe tips (05ndash)10ndash30(ndash38) mm long 02ndash04(ndash06) mm wide at unbranched parts yellowish orreddish to brownish orange usually paler yellowto yellowish orange at terminal portions of lobessurface often shiny lobes 150ndash200(ndash250) mmthick attached to substrate by medullary hyphaethat sometimes form rhizine-like structures of con-glutinated hyphae (Fig 5) upper cortical layercomposed of thick-walled irregularly arranged hy-phae 5ndash15 mm and with 15ndash20 mm thick luminaalgal layer composed of clustered trebouxoid algaemedulla rather compact lower cortex absent orpresent where lobes lift from substrate (Fig 6) pro-thallus occasionally present orange thin visiblebetween lobes and at thallus margin pseudocy-phellae sometimes present Apothecia mainly pre-sent in central part often abundant occasionallylacking 04ndash27 mm diam stipitate stipe 03ndash10mm disk concave to plane often flexuose orangeto reddish or brownish orange margin yellowishorange true exciple 50ndash60 mm thick in lateral por-

2002] 303KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 1 Habit of Caloplaca gyalectoides holotype (LD) Scale bars a and b 5 1 mm

tion composed of thick-walled irregular arrangedhyphae thalline exciple very thick 75ndash125 mmyellow to orange mostly paler than and raisedabove or level with disk sometimes with radiatingfurrows hymenium 80ndash100 mm high with yellow-brownish epithecium 5ndash12(ndash20) mm thick hypoth-ecium hyaline 25ndash80 mm thick paraphyses septatesimple or apically branched ca 15 mm thick in the

middle part apically slightly swollen to 3 mmdiam asci cylindric 50ndash80 3 10ndash12 mm asco-spores narrowly ellipsoid (12ndash)13ndash17(ndash19) 3(40ndash)45ndash60(ndash75) mm septum (30ndash)40ndash60(ndash75) mm thick sometimes curved spermogoniaprotruding reddish orange ca 01ndash03 mm widespermatia narrowly ellipsoid-bacilliform 33ndash50 309ndash12 mm Thallus and apothecia K1 red C2 I2

304 [VOL 105THE BRYOLOGIST

FIGURE 2 Known distribution of Caloplaca gyalec-toides

FIGURE 3 Bundles of hyphae on the thalline marginof Caloplaca gyalectoides Scale bar 5 10 mm

Chemistry Caloplaca chilensis contains theanthraquinones parietin and its oxidation productsteloschistin fallacinal parietinic acid and emodinParietin is the dominant compound in the apothecia(89) In the thallus parietin accounts for 55 ofthe anthraquinones and teloschistin fallacinal par-ietinic acid and emodin accounts for 15 264 and 1 respectively This anthraquinone che-mosyndrome agrees with A3 of Soslashchting (1997) Itmust be borne in mind that the calculated propor-tions are not absolute quantities but are based onabsorption at 270 nm

Ecology and distribution Caloplaca chilensisgrows on branches of bushes of various sizes aswell as on small trees and cacti The phorophytespecies does not seem to be important as it wasfound growing on at least six different phorophyteshowever always sun exposed The species occurson both dead and living twigs and branches It ismostly found in localities with dry open shrub veg-etation that is often grazed by sheep or other ani-mals (Fig 7) Even though boulders occur in mostknown localities of C chilensis and these oftenhold a large number of other Caloplaca species Cchilensis was never found on rocks The known lo-calities span from about 25 to 1100 m above sealevel However it is likely that C chilensis occursat even higher altitudes In most known localitiesC chilensis was abundant

A number of other species are often found to-gether with C chilensis eg Candelaria sp Chry-sothrix pavonii (Fr) J R Laundon C cf cerinaTeloschistes chrysophthalmus (L) Th Fr Ramal-ina sp Xanthoria ascendens S Kondratyuck andUsnea spp

Caloplaca chilensis apparently is restricted to thepart of Chile (Fig 8) that is characterized by heavyfogs caused by upwelling cold currents along the

coast The currents outside the coast of the Atacamaproduce a stable mild climate with very little pre-cipitation The lichens there receive their water al-most exclusively from the fogs The effect of thefogs on the lichen communities varies dependingon distance from the sea elevation and topogra-phy Normally the fog comes in from the sea dur-ing the night and disappears around noon the nextday (Rundel 1978) Most of the known localitiesfor the species are hills or mountains near the coastbut two localities differ The first is located in thebottom of an inland valley where the fog movesinland The second one is located on a mountain atthe bottom of this valley about 40 km from thecoast There are however no mountains farther tothe coast blocking the way for the fog which canmove inland in such valleys

The described conditions are typical for an areaalong the coast in Peru and the northern part ofChile It is therefore likely that C chilensis will befound to have a larger distributional area than re-ported here However it was not found around Ar-ica in northern Chile or in southern Peru possiblybecause the climate there is too dry Caloplaca chi-lensis does not match any species on the list ofspecies recorded from Chile (Galloway amp Quilhot2001) or any of the species recorded from the re-gion (Follmann 1962 1967 Follmann amp Redon1972 Rasanen 1936 Zahlbruckner 1925)

Taxonomic notes It is remarkable that such acommon and conspicuous species as C chilensisdoes not seem to have been described earlier or atleast reported under some other name as many li-chenologists have visited some of the known lo-calities eg P N Fray Jorge One possible reasoncould be that it has expanded its distribution rangeonly recently in connection with the drastic and insome cases catastrophic changes in the lichen floraof the coastal mountains that has occurred duringthe last decades (Follmann 1995)

Caloplaca chilensis is morphologically similar to

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

2002] 303KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 1 Habit of Caloplaca gyalectoides holotype (LD) Scale bars a and b 5 1 mm

tion composed of thick-walled irregular arrangedhyphae thalline exciple very thick 75ndash125 mmyellow to orange mostly paler than and raisedabove or level with disk sometimes with radiatingfurrows hymenium 80ndash100 mm high with yellow-brownish epithecium 5ndash12(ndash20) mm thick hypoth-ecium hyaline 25ndash80 mm thick paraphyses septatesimple or apically branched ca 15 mm thick in the

middle part apically slightly swollen to 3 mmdiam asci cylindric 50ndash80 3 10ndash12 mm asco-spores narrowly ellipsoid (12ndash)13ndash17(ndash19) 3(40ndash)45ndash60(ndash75) mm septum (30ndash)40ndash60(ndash75) mm thick sometimes curved spermogoniaprotruding reddish orange ca 01ndash03 mm widespermatia narrowly ellipsoid-bacilliform 33ndash50 309ndash12 mm Thallus and apothecia K1 red C2 I2

304 [VOL 105THE BRYOLOGIST

FIGURE 2 Known distribution of Caloplaca gyalec-toides

FIGURE 3 Bundles of hyphae on the thalline marginof Caloplaca gyalectoides Scale bar 5 10 mm

Chemistry Caloplaca chilensis contains theanthraquinones parietin and its oxidation productsteloschistin fallacinal parietinic acid and emodinParietin is the dominant compound in the apothecia(89) In the thallus parietin accounts for 55 ofthe anthraquinones and teloschistin fallacinal par-ietinic acid and emodin accounts for 15 264 and 1 respectively This anthraquinone che-mosyndrome agrees with A3 of Soslashchting (1997) Itmust be borne in mind that the calculated propor-tions are not absolute quantities but are based onabsorption at 270 nm

Ecology and distribution Caloplaca chilensisgrows on branches of bushes of various sizes aswell as on small trees and cacti The phorophytespecies does not seem to be important as it wasfound growing on at least six different phorophyteshowever always sun exposed The species occurson both dead and living twigs and branches It ismostly found in localities with dry open shrub veg-etation that is often grazed by sheep or other ani-mals (Fig 7) Even though boulders occur in mostknown localities of C chilensis and these oftenhold a large number of other Caloplaca species Cchilensis was never found on rocks The known lo-calities span from about 25 to 1100 m above sealevel However it is likely that C chilensis occursat even higher altitudes In most known localitiesC chilensis was abundant

A number of other species are often found to-gether with C chilensis eg Candelaria sp Chry-sothrix pavonii (Fr) J R Laundon C cf cerinaTeloschistes chrysophthalmus (L) Th Fr Ramal-ina sp Xanthoria ascendens S Kondratyuck andUsnea spp

Caloplaca chilensis apparently is restricted to thepart of Chile (Fig 8) that is characterized by heavyfogs caused by upwelling cold currents along the

coast The currents outside the coast of the Atacamaproduce a stable mild climate with very little pre-cipitation The lichens there receive their water al-most exclusively from the fogs The effect of thefogs on the lichen communities varies dependingon distance from the sea elevation and topogra-phy Normally the fog comes in from the sea dur-ing the night and disappears around noon the nextday (Rundel 1978) Most of the known localitiesfor the species are hills or mountains near the coastbut two localities differ The first is located in thebottom of an inland valley where the fog movesinland The second one is located on a mountain atthe bottom of this valley about 40 km from thecoast There are however no mountains farther tothe coast blocking the way for the fog which canmove inland in such valleys

The described conditions are typical for an areaalong the coast in Peru and the northern part ofChile It is therefore likely that C chilensis will befound to have a larger distributional area than re-ported here However it was not found around Ar-ica in northern Chile or in southern Peru possiblybecause the climate there is too dry Caloplaca chi-lensis does not match any species on the list ofspecies recorded from Chile (Galloway amp Quilhot2001) or any of the species recorded from the re-gion (Follmann 1962 1967 Follmann amp Redon1972 Rasanen 1936 Zahlbruckner 1925)

Taxonomic notes It is remarkable that such acommon and conspicuous species as C chilensisdoes not seem to have been described earlier or atleast reported under some other name as many li-chenologists have visited some of the known lo-calities eg P N Fray Jorge One possible reasoncould be that it has expanded its distribution rangeonly recently in connection with the drastic and insome cases catastrophic changes in the lichen floraof the coastal mountains that has occurred duringthe last decades (Follmann 1995)

Caloplaca chilensis is morphologically similar to

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

304 [VOL 105THE BRYOLOGIST

FIGURE 2 Known distribution of Caloplaca gyalec-toides

FIGURE 3 Bundles of hyphae on the thalline marginof Caloplaca gyalectoides Scale bar 5 10 mm

Chemistry Caloplaca chilensis contains theanthraquinones parietin and its oxidation productsteloschistin fallacinal parietinic acid and emodinParietin is the dominant compound in the apothecia(89) In the thallus parietin accounts for 55 ofthe anthraquinones and teloschistin fallacinal par-ietinic acid and emodin accounts for 15 264 and 1 respectively This anthraquinone che-mosyndrome agrees with A3 of Soslashchting (1997) Itmust be borne in mind that the calculated propor-tions are not absolute quantities but are based onabsorption at 270 nm

Ecology and distribution Caloplaca chilensisgrows on branches of bushes of various sizes aswell as on small trees and cacti The phorophytespecies does not seem to be important as it wasfound growing on at least six different phorophyteshowever always sun exposed The species occurson both dead and living twigs and branches It ismostly found in localities with dry open shrub veg-etation that is often grazed by sheep or other ani-mals (Fig 7) Even though boulders occur in mostknown localities of C chilensis and these oftenhold a large number of other Caloplaca species Cchilensis was never found on rocks The known lo-calities span from about 25 to 1100 m above sealevel However it is likely that C chilensis occursat even higher altitudes In most known localitiesC chilensis was abundant

A number of other species are often found to-gether with C chilensis eg Candelaria sp Chry-sothrix pavonii (Fr) J R Laundon C cf cerinaTeloschistes chrysophthalmus (L) Th Fr Ramal-ina sp Xanthoria ascendens S Kondratyuck andUsnea spp

Caloplaca chilensis apparently is restricted to thepart of Chile (Fig 8) that is characterized by heavyfogs caused by upwelling cold currents along the

coast The currents outside the coast of the Atacamaproduce a stable mild climate with very little pre-cipitation The lichens there receive their water al-most exclusively from the fogs The effect of thefogs on the lichen communities varies dependingon distance from the sea elevation and topogra-phy Normally the fog comes in from the sea dur-ing the night and disappears around noon the nextday (Rundel 1978) Most of the known localitiesfor the species are hills or mountains near the coastbut two localities differ The first is located in thebottom of an inland valley where the fog movesinland The second one is located on a mountain atthe bottom of this valley about 40 km from thecoast There are however no mountains farther tothe coast blocking the way for the fog which canmove inland in such valleys

The described conditions are typical for an areaalong the coast in Peru and the northern part ofChile It is therefore likely that C chilensis will befound to have a larger distributional area than re-ported here However it was not found around Ar-ica in northern Chile or in southern Peru possiblybecause the climate there is too dry Caloplaca chi-lensis does not match any species on the list ofspecies recorded from Chile (Galloway amp Quilhot2001) or any of the species recorded from the re-gion (Follmann 1962 1967 Follmann amp Redon1972 Rasanen 1936 Zahlbruckner 1925)

Taxonomic notes It is remarkable that such acommon and conspicuous species as C chilensisdoes not seem to have been described earlier or atleast reported under some other name as many li-chenologists have visited some of the known lo-calities eg P N Fray Jorge One possible reasoncould be that it has expanded its distribution rangeonly recently in connection with the drastic and insome cases catastrophic changes in the lichen floraof the coastal mountains that has occurred duringthe last decades (Follmann 1995)

Caloplaca chilensis is morphologically similar to

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

2002] 305KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 4 Habit of Caloplaca chilensis holotype Froden amp Arup 1 (LD) Scale bar 5 1 mm

FIGURE 5 Rhizine-like structures on Caloplaca chi-lensis holotype Froden amp Arup 1 (LD) Scale bar 5 01mm

FIGURE 6 Transverse section of lobe Caloplaca chi-lensis showing presence of lower cortical layer composedof thick-walled irregulary arranged hyphae Skottsberg1955 (S) Scale bar 5 10 mm

many other lobate species of the genus eg thesaxicolous species C biatorina (A Massal) JSteiner and C ignea Arup or even the corticolousXanthoria tenax L Lindblom that occurs in similarhabitats in southwestern North America It can beseparated from these by the structure of the cortexthat is composed of thick-walled irregularly ar-ranged hyphae with long and narrow lumina not aparaplectechyma The lobe tips of X tenax are flat-tened in contrast to the ones in C chilensis that areconvex in all parts The cortex structure of C chi-lensis is similar to the pseudoprosenchyma definedby Kondratyuk and Karnefelt (1997) but the hy-phae differ in being less gelatinized and congluti-nated This structure may also be found in the sax-

icolous species C brattiae W A Weber C ver-ruculifera (Vainio) Zahlbr C lucens (Nyl) ZahlbrC trachyphylla (Tuck) Zahlbr and C coralloides(Tuck) Hult All species except C brattiae and Ccoralloides are larger than C chilensis and in ad-dition C verruculifera is isidiate Caloplaca brat-tiae differs mainly by the much smaller spores 97ndash136 mm and C coralloides differs by the mainlysubfruticose thallus with terete lobes

Generic affinity Caloplaca chilensis holds an

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

306 [VOL 105THE BRYOLOGIST

FIGURE 7 Typical habitat for Caloplaca chilensis incoastal Chile ca 20 km N of Huasco Photo P Froden

intermediate position between Caloplaca and Xan-thoria as far as the growth form is concerned How-ever true species of Xanthoria have paraplecten-chymatous cortical layers whereas C chilensishave prosoplectenchyma (Fig 6) Various corticaltissues are however represented in the much moreheterogeneous Caloplaca The position outsideXanthoria is further supported by molecular evi-dence as an ITS sequence (data not presented)proved to have no similarity to those of Xanthoriaor closely related placodioid species The DNAdata clearly showed that C chilensis belongs inCaloplaca as presently defined but no closely re-lated species could be identified Further analysesincluding other genes and more species are neces-sary to find its closest relatives

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup19 20 21 22 23 S of Freirina ca 16 km along roadfrom Freirina to Quebradita just S of Questa la Totora28836919 S 71806739 W Froden amp Arup 24 25 26 2829 30 31 32 Region IV La Serena ca 25 km N of LaSerena close to Punta Poroto just E of Routa 5 2984529S 71819829 W Froden amp Arup 8 9 La Serena ca 15km N of La Serena Punta Teatinos 29849599 S71817649 W Froden amp Arup 10 11 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-Quedabra

Seca 30832839 S 71827319 W Froden amp Arup 12 13Ovalle Fray Jorge NP at entrance of park 30838569 S71836799 W Froden amp Arup 34 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 16 17 La Hig-uera road between Mina de Tofo and Chungungo W sideof mountain 29827409 S 71816459 W Froden amp Arup33 (all specimens located at LD) Ovalle Las Cardas 1V1955 Skottsberg (S)

ARTHONIA TETRASPORA S Kondratyuk amp Karne-felt sp nov FIG 9

Ab omnibus Arthoniis lichencolis differt ascis tetras-poribus tantum et sporis 1ndash2(ndash3)-septatis

TYPE CHILE Dept Ovalle Las Cardas on Oxalis gi-gantea on thalli and apothecia of Caloplaca chilensis1v1955 CA Skottsberg (UPS holotype GZU KW LD SGOisotypes)

On thallus and thalline margin of apothecia ofhost lichen Caloplaca chilensis forming black ir-regular to rounded ascomata that become plane orconvex primarily immersed in host tissue 03ndash05mm across and 02ndash03 mm high (Fig 8) hypoth-ecium blackish-brown 375ndash700(ndash2000) mmthick K1 becoming somewhat blackish withgreenish tinge epihymenium ca 12 mm thickblack-brown with greenish shade K2 or somewhatbrownish black hymenium 40ndash48(ndash60) mm highJ1 blue paraphysoides up to 23 mm thick withbrown-blackish cups and slightly widened tips 28ndash32 mm thick asci broadly clavate (285ndash)310ndash335(ndash380) 3 (17ndash)18ndash19 mm with reddish-orangeor brownish-orange pigment with 4 spores asco-spores (1ndash)2(ndash3)-septate clavate with middle cellnarrower than others at first hyaline later becom-ing brownish with pigment grains (14ndash)15ndash18 360ndash75 mm sometimes with clearly visible epis-porium spermogonia at basis of ascomata sper-matia bacilliform ca 4 3 1 mm

Ecology and distribution The habitat ecologyand distribution of A tetraspora is almost identicalwith that of its host C chilensis (Fig 10)

Taxonomic notes Arthonia tetraspora differsfrom all other known lichenicolous species in thegenus in having the only 4-spored asci as well asby the (1ndash)2(ndash3)-septate ascospores According tothe number of spores it belongs to the A destruensRehm ap Arnold complex that is known mainly asparasites on species of the genus Physcia (P ai-polia P stellaris and P tenella) and on Xanthoriaparietina in Europe However A tetraspora differsfrom A destruens by the 4-spored asci while Adestruens has 2ndash4ndash8-spored asci Furthermore Atetraspora has (1ndash)2(ndash3) septate and slightly largerascospores and narrower paraphysoides while Adestruens only has 2-septate spores that are slightlysmaller 10ndash16 3 3ndash6 mm

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

2002] 307KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

FIGURE 8 Known distribution of Caloplaca chilensis

FIGURE 9 Habit of Arthonia tetraspora Froden ampArup 2 (LD) Scale bar 5 01 mm

The new species can be confused with some lich-enicolous species of the genus Opegrapha Ope-grapha physciaria (Nyl) D Hawksw amp Coppinsknown from Xanthoria and Caloplaca is howevercharacterized by 3-septate spores and 8-spored asciin addition to several other important morphologi-cal and chemical characters Furthermore specieswith 4-spored asci in Opegrapha (O brevis Cop-pins and O pertusariicola Coppins amp P James)grow on representatives of the genus Thelotremaand have 3-septate or (5ndash)6-septate ascospores

ParatypesmdashCHILE Region III Road to Carizzal Bajofrom Huasco a few km N of Agua Luna and ca 20 kmN of Huasco 28818339 S 71809379 W Froden amp Arup18 Region IV La Serena ca 25 km N of La Serena closeto Punta Poroto just E of Routa 5 2984529 S 71819829W Froden amp Arup 2 3 4 5 6 7 Ovalle along roadOvalle-Recoleta between Cerillos de Taymar-QuedabraSeca 30832839 S 71827319 W Froden amp Arup 14 Oval-le Fray Jorge NP at the entrance of park 30838569 S71836799 W Froden amp Arup 35 Illapel along road be-tween Illapel-Salamanca a few km SE of Illapel31841929 S 71809219 W Froden amp Arup 15 (all speci-mens at LD)

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

308 [VOL 105THE BRYOLOGIST

FIGURE 10 Known distribution of Arthonia tetraspora

ACKNOWLEDGMENTS

IK received financial support from Crafoordska Stiftel-sen and Magn Bergvalls Stiftelse SK received financialsupport from INTAS (ref No 97-30778) and US receivedfinancial support from the Carlsberg Foundation PF andUA received financial support for their fieldwork fromKungliga Fysiografiska sallskapet Kungliga Vetenskap-sakademien Stiftelsen Lars Hiertas minne Ekedahl-Lund-bergska fonden Ove Almborns fond D amp E Ekvallsfond and Per Westlings minnesfond Their contributionsare very much acknowledged Lene Christiansen operatedthe HPLC-equipment Else Meyer Andersen and CharlotteHansen assisted with the DNA laboratory work Per Las-sen translated the diagnoses to Latin and Susanna Riebeassisted with the photographs They are all thanked fortheir assistance

LITERATURE CITED

ARUP U 1995 Littoral species of Caloplaca in NorthAmerica A summary and key THE BRYOLOGIST 98129ndash140

CLAUZADE G amp C ROUX 1985 Likenoj de okcidentaEuropo Ilustrita Determinlibro Royan

DODGE C W 1971 Some lichens of tropical Africa VLecanoraceae to Physciaceae Beiheft zur Nova Hed-wigia 38 1ndash225

FEUERER T 2001 Checklist of South African lichens andlichenicolous fungi Version 1 April 2001 http

wwwun i -hamburg de b io log ie i a lb he rba r souafpf2htm

FOLLMANN G 1962 Eine borkenbewohnende Flechten-geschellschaft der zentralchilensischen Dornstrauch-formationen mit kennzeichnendem Teloschistes chry-sophthalmus Nova Hedwigia 4 109ndash129

mdashmdashmdash 1967 Die Flechtenflora der nordchilenischen Ne-beloase Cerro Moreno Nova Hedwigia 14 263ndash267

mdashmdashmdash 1995 On the impoverishment of the lichen floraand the retrogression of the lichen vegetation in coast-al central and northern Chile during the last decadesCryptogamic Botany 5 224ndash231

mdashmdashmdash amp J REDON 1972 Erganzungen zur Flechtenflorader nordchilenischen Nebeloasen Fray Jorge und Tal-inay Willdenowia 6 431ndash460

GALLOWAY D J 1985 Flora of New Zealand LichensP D Hasselburg Government Printer Wellington

mdashmdashmdash amp W QUILHOT 2001 Checklist of Chilean lichensand lichenicolous fungi Version 1 March 2001 httpwwwuni -hamburg de b io logie ia lb herbar chilepf2htm

HANSEN E J POELT amp U SoslashCHTING 1987 Die Flechten-gattung Caloplaca in Gronland Meddelelser omGroslashnland Bioscience 25 1ndash52

KARNEFELT I 1989 Morphology and phylogeny in theTeloschistales Cryptogamic Botany 1 147ndash203

mdashmdashmdash S KONDRATYUK U SoslashCHTING amp P FRODEN 2002Xanthoria karrooensis and Xalexanderbaai (Telos-chistaceae) two new remarkable lichens from SouthernAfrica Lichenologist (in press)

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002

2002] 309KARNEFELT ET AL CALOPLACA FROM SOUTHERN HEMISPHERE

KONDRATYUK S amp I KARNEFELT 1997 Josefpoeltia andXanthomendoza two new genera in the Teloschista-ceae (Lichenized Ascomycotina) Bibliotheca Lichen-ologica 68 19ndash44

NAVARRO-ROSSINES P amp N HLADUN 1996 Las especiessaxicolo-calcicolas grupo de Caloplaca lactea (Telos-chistaceae liquenes) en las regiones mediterranea ymedioeuropea Bulletin de la Societe Linneenne deProvence 47 139ndash166

POELT J amp E HINTEREGGER 1993 Beitrage zur Kenntnisder Flechtenflora des Himalaya VII Die GattungenCaloplacaFulgensia und Ioplaca Bibliotheca Lich-enologica 50 1ndash265

PURVIS W B COPPINS D HAWKSWORTH P JAMES amp DMOORE 1992 The Lichen Flora of Great Britain andIreland British Lichen Society London

RASANEN V 1936 Collationes ad lichenologiam Chilen-sem pertinentes Revista Universitaria 21 137ndash148

RUNDEL P H 1978 Ecological relationships of desert fogzone lichens THE BRYOLOGIST 81 277ndash293

SoslashCHTING U 1997 Two major anthraquinone chemosyn-

dromes in Teloschistaceae pp 135ndash144 In R Turk ampR Zorer (eds) Progress and Problems in Lichenologyin the Nineties IAL 3 Bibliotheca Lichenologica 68

mdashmdashmdash 2001 Chemosyndromes with chlorinated anthra-quinones in the lichen genus Caloplaca pp 395ndash404In P M McCarthy G Kantvilas amp S H J J Lou-whoff (eds) Lichenological Contributions in Honourof Jack Elix Bibliotheca Lichenologica 78

WETMORE C M 2001 The Caloplaca citrina group inNorth and Central America THE BRYOLOGIST 104 1ndash11

mdashmdashmdash amp E I KARNEFELT 1998 The lobate and subfru-ticose species of Caloplaca in North and CentralAmerica THE BRYOLOGIST 101 230ndash255

mdashmdashmdash amp mdashmdashmdash 2000 What is Caloplaca cinnabarinaTHE BRYOLOGIST 102 683ndash691

ZAHLBRUCKNER A 1925 Chilenische Flechten gesam-melt von CA Skottsberg Meddelanden fran Gote-borgs botaniska Tradgard 2 1ndash26

ms received March 8 2002 accepted April 15 2002