Insights into the genus Austrella Pannariaceae Peltigerales),Key words: biogeography, Gondwanan...

The Lichenologist 49(1): 57–65 (2017) © British Lichen Society, 2017doi:10.1017/S0024282916000621

Insights into the genus Austrella (Pannariaceae, Peltigerales),including a new species from the Falkland Islands

Alan M. FRYDAY, Damien ERTZ and Per Magnus JØRGENSEN

Abstract:The new speciesAustrella isidioidea, which is unique in the genus in having isidioid structureson the thallus lobe ends as well as apothecia lacking a thalline margin, is described from the FalklandIslands. A collection with an identical mtSSU rDNA sequence to A. arachnoidea but with significantmorphological differences (viz. a variable apothecial margin ranging from an arachnoid hyphal weft to acorticated regular margin) is reported from Îles Kerguelen. The phylogenetic and biogeographicalimplications of these new records are discussed.

Key words: biogeography, Gondwanan distribution, Îles Kerguelen, lichen, Psoroma-clade

Accepted for publication 11 August 2016

Introduction

The small Australasian genus Austrella P. M.Jørg. is characterized within the Pannariaceaeby its elongated, thin-walled asci that lackamyloid apical structures and by young apo-thecia being surrounded by an arachnoidhyphal weft that disappears gradually with age,as observed in A. brunnea (P. M. Jørg. ) P. M.Jørg. (Jørgensen 2004). However, recentcollections referable to Austrella considerablyextend the morphological variation andgeographical range of the genus and theimplications of these are discussed here.

Materials and Methods

The study is based on specimens collected during recentfieldwork on the Falkland Islands and Îles Kerguelen bythe first and second authors, respectively. Apothecialcharacteristics were examined by light microscopy onhand-cut sections mounted in water. Thallus sections

were investigated, and anatomical measurements made,in 10% KOH. Standard chemicals were used for spot-test reactions and thin-layer chromatography followedthe methods of Orange et al. (2001).

Molecular techniques

Hand-prepared sections of the hymenium were usedfor direct PCR as described in Ertz et al. (2015). Theouter wall of ascomata was removed with a sterile razorblade to isolate the hymenium that was rinsed withacetone. The material was then added to a tube con-taining the PCR reaction mixture and amplified directly.A targeted fragment of c. 0·8 kb of the mtSSU rDNAwasamplified using primers mrSSU1 and mrSSU3R (Zolleret al. 1999). Both strands were sequenced byMacrogen®

using amplification primers. Sequence fragments wereassembled with Sequencher version 4.6 (Gene CodesCorporation, Ann Arbor, Michigan). Sequences weresubjected to MegaBLAST searches (Zhang et al. 2000;Wheeler et al. 2006) to verify their closest relatives and todetect potential contaminations.

Phylogenetic analyses

The mtSSU matrix published by Ekman et al. (2014)and deposited under study number 14978 in the Tree-BASE repository (http://www.treebase.org) was used as atemplate to include our two newly sequenced Austrellasamples. The sequences were aligned manually usingMesquite v.3.04 (Maddison & Maddison 2015). Theresulting mtSSU matrix consisted of 98 sequences and726 unambiguously aligned characters, of which 344were variable.

The best-fit model of DNA evolution TIM+I+Gwas chosen using the Akaike information criterion(AIC; Akaike 1973) as implemented in Modeltest v. 3.7(Posada & Crandall 1998). Bayesian analyses were

A. M. Fryday: Herbarium, Department of PlantBiology, Michigan State University, East Lansing, MI48824-1312, USA. Email: [email protected]. Ertz: Department Bryophytes-Thallophytes, BotanicGarden Meise, Nieuwelaan 38, B-1860 Meise, Belgium;and Fédération Wallonie-Bruxelles, Direction Généralede l’Enseignement non obligatoire et de la RechercheScientifique, Rue A. Lavallée 1, B-1080 Bruxelles,Belgium.P. M. Jørgensen: Herbarium, Museum of NaturalHistory, Bergen University Museum, Allégaten 41,Postboks 7800, N-5020 Bergen, Norway.

at https:/www.cambridge.org/core/terms. https://doi.org/10.1017/S0024282916000621Downloaded from https:/www.cambridge.org/core. Michigan State University Libraries, on 20 Jan 2017 at 14:18:23, subject to the Cambridge Core terms of use, available

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carried out using the Metropolis-coupled Markov chainMonte Carlo method (MCMCMC) in MrBayes v. 3.2.6(Huelsenbeck & Ronquist 2001; Ronquist &Huelsenbeck 2003) on the CIPRES portal (Miller et al.2010). Analyses were run under the selected model ofnucleotide substitution with six rate categories. Twoparallel MCMCMC runs were performed, with each runusing four independent chains and 100 000 000 gen-erations, and sampling trees every 1000th generation.Convergence diagnostics were estimated using the PSRF(potential scale reduction factor) where values closer to 1indicated convergence between runs (Gelman &Rubin 1992), and using Tracer v. 1.6 by plotting thelog-likelihood values of the sample points againstgeneration time (Rambaut & Drummond 2007).Posterior probabilities (PP) were determined by calcu-lating a majority-rule consensus tree generated from the150 002 post burn-in trees of the 200 002 trees sampledby the two MCMCMC runs using the sumt option ofMrBayes. In addition, a Maximum Likelihood (ML)analysis was performed on the mtSSU data set usingGARLI (Zwickl 2006, v. 0.951 for OSX) with defaultsettings and a single most-likely tree was produced(lnL = − 6789·802089). We used 1000 bootstrappseudoreplicates to calculate a majority-rule consensustree in PAUP* 4.0b10 (Swofford 2002) to assess theMaximum Likelihood bootstrap values (ML-bs).ML-bs≥70% and PP≥95% were considered to besignificant. Phylogenetic trees were visualized usingFigTree v. 1.4.2 (Rambaut 2012).

Additional collections examined: Austrella cf. arachnoideaP. M. Jørg. Îles Kerguelen: Île Guillou, 49°28'57''S,69°49'06''E, 14m, feldfield, 2014, D. Ertz 18847(BG, BR, MSC).

Results

We obtained two new mtSSU sequencesbelonging respectively to the new speciesAustrella isidioidea andAustrella cf. arachnoidea(GenBank Accession numbers KY069281and KY069280). Unfortunately, we wereunable to obtain sequences forRPB1, the onlyother gene for which a sequence for Austrellais available in GenBank. The Bayesian treewe recovered did not contradict the ML treetopology for the strongly supported branchesand hence only the ML tree is shown (Fig. 1).The new species Austrella isidioidea clusters

with Santessoniella arctophila in a clade thatincludes Psoroma s. str. but withoutsupport, whereas A. cf. arachnoidea forms aseparate branch with A. arachnoidea in apolytomy including both the Psoroma s. str.clade and another containing Psorophorus andArctoparmeliella. Austrella cf. arachnoideaand A. isidioidea are nested within ‘Clade 2c’of Ekman et al. (2014), which corresponds tothe clade from Nebularia incrassata to Psoromahypnorum III in our mtSSU tree (Fig. 1).Branch support within this clade is poor,rendering generic delimitations difficult. Thiswas already highlighted by Ekman et al. (2014)despite the use of a three gene phylogeny.

Taxonomy

Austrella isidioidea P. M. Jørg. &Fryday sp. nov.

MycoBank No.: MB 817913

Austrellae brunneae similis, sed thallo atrofuscosquamulis adscententibus in parte isidoideis.

Typus: Falkland Islands, West Falkland, PortHoward, Mt. Maria, Lightning Rocks, 51·619028°S,59·601849°W, 575m., over Andreaea sp. in recess onexposed E-facing rocks, 26 January 2015, A. M. Fryday10918 (MSC—holotypus).

(Figs. 2–4)

Thallus forming cushions up to 3 cm diam.,composed of brown to dark brown, partlyascending squamules with a paler margin,becoming blackened at the apices and at spotsalong the edges that become raised and evolveinto isidioid structures 0·1–0·2mm thick and0·6mm tall; squamules finally becomingcompletely blackened (Fig. 2A & B).Cortex composed of a single layer ofthin-walled cells. Photobiont Nostoc, in shortchains enclosed in a reticulum of hyphae,individual cells 5–6 µm diam.

FIG. 1. Maximum Likelihood tree of Pannariaceae using GARLI and based on a data set of mtSSU sequences.Vahliella leucophaea in the Vahliellaceae is used as the rooting taxon. Internal branches with Maximum Likelihoodbootstrap values ≥70% obtained from a GARLI analysis are considered strongly supported and shown above internalbranches. Posterior probabilities ≥0·95 resulting from a Bayesian analysis are shown below internal branches. Internalbranches strongly supported by both analyses are represented by thicker lines. The new species is in bold. Collectingnumbers of the authors following the species names act as specimen and sequence identifiers. For clarity, support

values are not shown for some short internal branches of groups not related to Austrella.

58 THE LICHENOLOGIST Vol. 49


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0.03

P. microphyllizans

Fuscopannaria laceratula

P. hypnorum IV

Psorophorus pholidotus

F. pacifica I

L. mariana

Psoroma hypnorum III

P. rubiginella II

L. finmarkicum

Psoroma polychidioides

Nevesia sampaiana II

Degeliella versicolor

P. aff. immixta

F. pacifica II

P. atlantica III

Fuscopannaria ignobilis

P. plumbea I

Parmeliella triptophylla I

Pectenia plumbea II

P. athroophylla II

Ramalodium succulentum

P. implexa

Santessoniella arctophila

P. rubiginella I

P. isabellina

P. paleaceum

D. durietzii

Fuscoderma amphibolum

J. cephalodina I

P. dichroum

P. contorta

Degeliella rosulata

Leciophysma furfurascens

P. sphinctrina

P. fuegiensis

Protopannaria pezizoides II

P. hypnorum II

Nebularia incrassataF. applanatum I

P. multifida

P. pezizoides I

P. miradorensis

Moelleropsis nebulosa I

Physma byrsaeum

D. gayana III

P. insularis

P. cyanoloma

E. verruculosum

M. nebulosa II

Pannaria calophylla

Staurolemma oculatumS. omphalarioides

P. pannosa

P. radians

P. farinosa

F. sorediataF. praetermissa

L. pycnophorum

A. cf. arachnoidea 18847

N. sampaiana I

P. hookeri

P. nigrocincta

A. arachnoidea

Xanthopsoroma contextum

P. athroophylla I

P. saccharatum

D. duplomarginata

Fuscopannaria ahlneri

P. atlantica II

F. olivacea

Joergensenia cephalodina II

P. plumbea III

P. rubiginosa

P. plumbea IV

P. lurida ssp lurida

P. hypnorum V

Degelia gayana I

P. atlantica I

P. aphthosum

F. leucostictoides

F. leucosticta

Austrella isidioidea 10918 holotype

Vahliella leucophaea

Austroparmeliella lacerata

P. pallida

X. soccatum

F. maritima

Santessoniella sp.

Lepidocollema stylophora

Pannaria hispidula

P. lurida ssp russellii

P. appalachensis

Erioderma pedicellatum

P. hypnorum I

Siphulastrum squamosum

Leioderma erythrocarpum

P. triptophylla II

M. nebulosa III

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1AUSTRELLA

2017 Austrella—Fryday et al. 59



Apothecia fairly abundant but difficult toobserve among the isidia, sessile, partlyclustered, black, ≤2mm diam. with distinctproper exciple, ≤200 µm wide (Fig. 2D).Hymenium non-amyloid, 120–150 µmhigh ofsimple, septate hyphae, apically with darkpigment. Asci to 150 µm, slender, thin-walled, I+ blue, but without apical amyloidstructures (Fig. 2C); ascospores hyaline,ellipsoid, 17–19(20)× 8–9(10) µm often withoil drops and occasionally appearing septate(pseudoseptate) (Fig. 2E).Conidiomata not observed.

Chemistry. No lichen substances detected(by TLC).

Remarks. Austrella isidioidea is as yet knownfrom only two collections from the FalklandIslands (Figs. 3 & 4). At its type locality, itgrew amongmosses on lichen-rich rock ledgesin association with Shackletonia siphonospora(Olech & Søchting) Søchting et al., Endocenainformis Cromb., Rimularia andreaeicolaFryday and Thamnolia vermicularis (Sw.)Schaer., whereas adjacent rocks supportedHimantormia deusta (Hook. f.) A. Thell &Søchting, Ochrolechia antarctica (Müll. Arg.)Darb., Pertusaria macloviana Müll. Arg.,Poeltidea perusta (Nyl.) Hertel &Hafellner, andUsnea aurantiacoatra (Jacq.) Bory (Fig. 4). Atits only other known locality (Fryday 11427;see below for details) it occurred in a similar

A

B

C

D E

FIG. 2. Austrella isidioidea. A, thallus; B, isidia; C, ascus; D, apothecia; E, ascospore. Note the pseudoseptum in theascospore and the absence of apical amyloid structures in the ascus (A, Fryday 11427; B–E, Fryday 10918,

holotype). Scales: A & D = 1mm; B = 0·5mm; C = 25 µm; E = 10 µm. In colour online.




51º S 61º W 60º W 59º W 58º W

51.5º S

52º S

52.5º S

N

FIG. 3. Known localities for A. isidioidea on the Falkland Islands.

FIG. 4. Type locality of Austrella isidioidea showing the lichen-rich rock ledges with Himanthormia lugubris(dark thalli) and several crusts including the large, conspicuous white patches of Ochrolechia antarctica.

In colour online.




situation to the type collection amongAndreaea sp. where it was associated withBartlettiella fragilisD. J. Galloway& P.M. Jørg.The habit of the lichen, a dark basal thallusmore or less hidden among the moss withwhich it grows and the squamules extendinginto isidia that grow up through the moss,make A. isidioidea hard to observe in the fieldand it is probably quite frequent in suitablehabitats and may occur elsewhere in southernSouth America.The gross morphology of the new species

gives it a superficial resemblance to somespecies of Siphulastrum Müll. Arg., a genusdescribed from the nearby Isla de losEstados, and also frequent on the FalklandIslands. However, the ‘isidia’ of Siphulastrumare densely compacted together formingan almost continuous crust, whereas inA. isidioidea they are more widely dispersedand separated. More critically, the apotheciaof A. isidioidea clearly differ from those ofSiphulastrum by the asci lacking amyloidapical structures, as well as the thallus lackinglichen substances (argopsin in Siphulastrum)and having thin-walled cortical cells.Morphologically,Austrella isidioidea appears

to be closest to A. brunnea (P. M. Jørg.) P. M.Jørg. from New Zealand, a species also form-ing cushions among bryophytes on theground, but with flat, brown squamules andsmaller, consistently non-septate ascospores.

Additional specimen examined. Falkland Islands: EastFalkland: Mt. Usborne, W-facing outcrop just belowsummit, 51·691324°S, 58·839517°W, 610m., overAndreaea sp. on rock ledge, 2015, Fryday 11427 (MSC).

Discussion

Systematic position of Austrellaisidioidea

In the molecular analysis of Ekman et al.(2014), Austrella arachnoidea (the typespecies of the genus) grouped with severalother small genera or species complexes inthe difficult psoromoid section (Clade 2c).Relationships in this clade, which roughlycorresponds to small species assigned to thegenus Psoroma, are currently unclear and inneed of further investigation.

Our molecular analysis clearly places the newspecies in Clade 2c of Ekman et al. (2014)although it did not form a monophyletic groupwith A. arachnoidea (Fig. 1). As A. isidioideashares the distinctive ascus structure ofA. arachnoidea described above, this clearlyrepresents a challenge in deciding in whichgenus to place the new species: a widelycircumscribed Psoroma as suggested bymolecular results, or Austrella as suggested bymorphology.Of the three genes used by Ekman et al.

(2014), only sequences for the small subunitof the mitochondrial ribosomal RNA(mtSSU) and the largest subunit of RNApolymerase II (RPB1) genes were available inGenBank for A. arachnoidea. Unfortunately,we were unable to obtain sequences forRPB1, so our phylogeny is based only on themtSSU gene and, consequently, the relativepositions of the two Austrella species in ourphylogenetic tree are based on a single sharedgene and this, combined with the low supportof the tree, places considerable doubt on therelationship revealed. We considered a con-strained monophyly test but concluded that,based on such a poorly resolved clade anddata, this was unlikely to solve the problem ofgeneric delimitation.Therefore, we decided to focus on mor-

phological evidence and as our new species isclearly not congeneric with the type specimenof Psoroma (P. hypnorum (Vahl) Gray), and iscloser to the type ofAustrella (A. arachnoidea),describing it in Austrella was considered thebest course of action.

Morphological variation in Austrellaarachnoidea

In addition to the molecular uncertaintiessurrounding the systematic position ofAustrella, detailed examination also revealedinconsistencies in the morphological char-acters that have been used to characterizethe genus. A collection from Îles Kerguelen(Ertz 18847; see Materials and Methodsfor details) had an identical mtSSU sequenceto A. arachnoidea (unfortunately, the ITS ofthe latter was not available fromGenBank forfurther comparison) but differed from it in




gross morphology in the apothecia having adistinct, regular thalline margin (Fig. 5A),and also anatomically in some asci havinga distinct KI+ blue ring structure in theirapex similar to that in Psoroma (Fig. 6).However, immature apothecia on theunderside of the specimen (i.e. in a shadedsituation) did possess an arachnoid margin(Fig. 5B) and also had asci lacking any KI+blue apical apparatus. The sequence datawere obtained from an apothecium with athick thalline margin. The apothecia ofAustrella isidioidea were also anomalous incompletely lacking an arachnoid margin,even when immature. However, the asci weretypical for those of the genus, being elon-gated and thin-walled and completely lackingany KI+ blue apical apparatus (Fig. 2C).

Biogeographical implications

Species of the genus Austrella previouslyhad a relatively restricted distribution, being

known only from the South Island ofNew Zealand and SE Australia (Jørgensen2004; Fig.7). The new records described hereextend the distribution to other parts ofthe southern cool temperate/subantarcticzone giving the genus a typical Gondwanandistribution. This highly disjunctive patternis well known among lichens in general, andgenera of the Pannariaceae in particular (e.g.Siphulastrum, Psoromaria, Psorophorus; cf.Galloway 1987). However, as shown byFryday & Coppins (2007), the Gondwanadistribution pattern can be further refined intooceanic and continental Gondwanan ele-ments with the oceanic element beingrestricted to SW Chile and the islands ofthe New Zealand Campbell Plateau, and thecontinental element extending from southernChile, through Tierra del Fuego, the islandgroups of the southern Atlantic and IndianOceans (including the Falkland Islands andÎles Kerguelen) to SE Australia/Tasmaniaand New Zealand. The oceanic element is

A

B

FIG. 5. Austrella cf. arachnoidea (Ertz 18847).A, apothecia on upper side with thick thalline margin;B, apothecia on underside with arachnoid margin.

Scales: A & B = 0·5mm. In colour online.

FIG. 6. Austrella cf. arachnoidea (Ertz 18847), ascus.Scale = 10 µm. In colour online.




characterized by the presence of a number ofspecies (e.g. Lithographa olivacea Fryday,Pertusaria stellata Fryday, Steinera symptychia(Tuck.) T. Sprib. & Muggia) that are absentfrom other southern subpolar regions, and theabsence of other conspicuous species (e.g.Poeltidea perusta (Nyl.) Hertel & Hafellner,Usnea sect. Neuropogon spp.) that are commonin the continental element. The distribution ofAustrella fits perfectly the continental Gondwa-nan distribution pattern.

The first author thanks the UK Government for funding(via theDarwin Initiative) to visit the Falkland Islands andcollect lichens, and Falklands Conservation, in particularDafydd Crabtree for logistical support. The secondauthor acknowledges the Fonds National de la RechercheScientifique from Belgium (FNRS) for financial supportand the Institut Polaire Français Paul-Emile Victor (IPEV- program 136 SUBANTECO, in particular Dr DavidRenault and Dr Maryvonne Charrier) for logisticalsupport to visit and collect on Îles Kerguelen.

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FIG. 7. Known world distribution of the genus Austrella. Key: ■ A. arachnoidea; ▲ A. cf. arachnoidea;● A. brunnea;⋆ A. isidioidea.




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http://mesquiteproject.orghttp://tree.bio.ed.ac.uk/software/figtree/http://beast.bio.ed.ac.uk/https:/www.cambridge.org/core/termshttps://doi.org/10.1017/S0024282916000621https:/www.cambridge.org/core

Insights into the genus Austrella (Pannariaceae, Peltigerales), including a new species from the Falkland IslandsIntroductionMaterials and MethodsMolecular techniquesPhylogenetic analyses

ResultsTaxonomyAustrella isidioidea P. M. Jørg. & Fryday sp. nov

Fig. 1Maximum Likelihood tree of Pannariaceae using GARLI and based on a data set of mtSSU sequences. Vahliella leucophaea in the Vahliellaceae is used as the rooting taxon. Internal branches with Maximum Likelihood bootstrap values ≥70% obFig. 2Austrella isidioidea. A, thallus; B, isidia; C, ascus; D, apothecia; E, ascospore. Note the pseudoseptum in the ascospore and the absence of apical amyloid structures in the ascus (A, Fryday 11427; B–E, Fryday 10918, holotype). Scales: A Fig. 3Known localities for A. isidioidea on the Falkland IslandsFig. 4Type locality of Austrella isidioidea showing the lichen-rich rock ledges with Himanthormia lugubris (dark thalli) and several crusts including the large, conspicuous white patches of Ochrolechia antarctica. In colour onlineDiscussionSystematic position of Austrella isidioideaMorphological variation in Austrella arachnoideaBiogeographical implications

Fig. 5Austrella cf. arachnoidea (Ertz 18847). A, apothecia on upper side with thick thalline margin; B, apothecia on underside with arachnoid margin. Scales: A & B=0·5&znbsp;mm. In colour onlineFig. 6Austrella cf. arachnoidea (Ertz 18847), ascus. Scale=10&znbsp;µm. In colour onlineFig. 7Known world distribution of the genus Austrella. Key: ■ A. arachnoidea; ▲ A. cf. arachnoidea; ● A. brunnea; ⋆ A. isidioidea

Insights into the genus Austrella Pannariaceae Peltigerales),Key words: biogeography, Gondwanan...

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