PhytoKeys 5: 31–44, doi: 10.3897/mycokeys.5.4140
Molecular data support placement of Cameronia in Ostropomycetidae (Lecanoromycetes, Ascomycota)
H. Thorsten Lumbsch 1, Gintaras Kantvilas 2, Sittiporn Parnmen 1
1 Department of Botany, Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, USA
2 Tasmanian Herbarium, Private Bag 4, Hobart, Tasmania, Australia 7001

Corresponding author: Thorsten Lumbsch (

Academic editor: P. Divakar

received 17 October 2012 | accepted 26 November 2012 | Published 30 November 2012

(C) 2012 H. Thorsten Lumbsch. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

For reference, use of the paginated PDF or printed version of this article is recommended.


The phylogenetic position of the Tasmanian endemic genus Cameronia Kantvilasis studied using partial sequences of nuclear LSU and mitochondrial SSU ribosomal DNA. Monophyly of the genus is supported, as is its placement in Ostropomycetidae, although its position within this subclass remains uncertain. Given the lack of close relatives to Cameronia and its morphological differences compared to other families with perithecioid ascomata in Ostropomycetidae, the new family Cameroniaceae Kantvilas & Lumbsch is proposed.


Cameroniaceae, lichens, new family, Tasmania, taxonomy


The lichen flora of Tasmania has a remarkable number of unique species, as well as several genera that are unknown or very rarely found in other regions. Examples include the genera Jarmania Kantvilas (Kantvilas 1996), Meridianelia Kantvilas & Lumbsch (Kantvilas and Lumbsch 2009), Siphulella Kantvilas, Elix & P. James(Kantvilas et al. 1992), Tasmidella Kantvilas, Hafellner & Elix (Kantvilas et al. 1999), and several species of Cladia (Kantvilas and Elix 1987, 1999) andthelotremoid Graphidaceae (Kantvilas and Vezda 2000; Mangold et al. 2009). In general, endemism can be either the result of survival of relict taxa (palaeoendemism) or recent speciation events (neoendemism) (Brandley et al. 2010; Brooks et al. 2006; Goldberg et al. 2005; Janssen et al. 2008; Kier et al. 2009; Kraft et al. 2010; Lamoreux et al. 2006; Olson et al. 2001; Qian 2001). The reasons for the relatively large amount of endemic taxa in Tasmania are not well understood. In the genus Cladia, for example, molecular data are consistent with a recent speciation and suggest neoendemism (Lumbsch et al. 2010; Parnmen 2011), but for most endemic taxa there are currently insufficient data available to test whether they represent relict lineages or are the product of recent speciation events. In some cases, however, lichens that were believed to be endemic to Tasmania, were subsequently also discovered in New Zealand, e.g. Bunodophoron flaccidum (Wedin 1993; Wedin 2001).

Lichen taxa unique to Tasmania include the genus Cameronia (Kantvilas 2012), which was recently described with an unclear systematic position and placed tentatively in Ostropomycetidae. The genus includes two species that occur on siliceous rocks at high elevations. Although its thallus is superficially similar to that of a species of Lecanora or Pertusaria, the genus is readily distinguished by the presence of eumuriform ascospores in thick-walled, broadly obovate, hemiamyloid asci with a non-amyloid tholus, formed in a hamathecium consisting of richly branched, anastomosing paraphysoids. The ascomata are perithecioid. Secondary metabolites present in the genus include the 9-O-methylpannaric acid chemosyndrome and an unknown triphenyl.

Thick-walled asci having a hemiamyloid wall and non-amyloid tholus, anastomosing paraphysoids and muriform ascospores are all characters reminiscent of Arthoniales (Ertz and Tehler 2011; Grube 1998; Tehler 1990), but the perithecioid ascomata, chlorococcoid photobiont, and morphological details of the ascus differ from this order (Kantvilas 2012). Perithecioid ascomata and thick-walled asci in a hamathecium consisting of anastomosing paraphysoids are characteristic for Protothelenellaceae and Thelenellaceae in Ostropomycetidae (Fryday and Coppins 2004; Mayrhofer 1987a, b; Mayrhofer and Poelt 1985; Schmitt et al. 2005). However, these families differ in having cylindrical asci and, furthermore, Thelenellaceae lacks any amyloid reactions of the asci, whereas Protothelenellaceae have an amyloid tholus. Because phenotypic characters do not place Cameronia in any group unambiguously and the placement in Ostropomycetidae was tentative, we used freshly collected material of the two species of Cameronia to generate DNA sequences of two loci (mtSSU and nuLSU rDNA) to test the monophyly of Cameronia and its placement of Cameronia in Ostropomycetidae, and to identify the closest relatives of the genus and place it in a family.

Materials and methods
Taxon sampling and molecular methods

The study is based on fresh material collected by GK and deposited in the Tasmanian Herbarium (HO) and the Field Museum of Natural History (F), and on DNA sequences downloaded from Genbank. Sequences of Umbilicariaceae were included as outgroup since this family has been shown previously to be sister to Lecanoromycetidae+Ostropomycetidae (Lumbsch et al. 2007a; Miadlikowska et al. 2006; Spatafora et al. 2006; Wedin et al. 2005). Sequence data of the two species of Cameronia were assembled with sequences of mitochondrial small subunit (mtSSU) and nuclear LSU rDNA downloaded from Genbank (Table 1). Sample preparation, DNA isolation, PCR and direct sequencing were performed as described previously (Mangold et al. 2008; Rivas-Plata and Lumbsch 2011). Primers for amplification were: mr SSU1 (Zoller et al. 1999) and MSU7 (Zhou and Stanosz 2001) for mtSSU, and AL2R (Mangold et al. 2008) and nu-LSU-1125-3’ (= LR6) (Vilgalys and Hester 1990) for nuLSU rDNA. Sequence fragments obtained were assembled with SeqMan 4.03 (DNASTAR) and manually adjusted.

Table 1.

Sequences obtained from Genbank for the study. Family or generic group as in figure 1, largely following (Lumbsch and Huhndorf 2010). Newly obtained sequences are indicated in bold.

Species Family/generic group as in Fig. 1 nuLSU mtSSU
Acarosporina microspora Stictidaceae AY584643 AY584612
Agyrium rufum - EF81824 EF81821
Ainoa mooreana - AY212850 AY212828
Anzina carneonivea - AY212829 AY212851
Arctomia delicatula Arctomiaceae AY853307 AY853355
Arctomia teretiuscula Arctomiaceae DQ007346 DQ007349
Aspicilia caesiocinerea Megasporaceae DQ780303 DQ780271
Aspicilia cinerea Megasporaceae DQ780304 DQ780272
Aspicilia contorta Megasporaceae DQ986782 DQ986876
Aspicilia hispida Megasporaceae DQ780305 DQ780273
Baeomyces placophyllus - AY300878 AF356658
Baeomyces rufus - DQ871008 DQ871016
Belonia russula Gyalectaceae FJ941887 AY648888
Bryophagus gloeocapsa Gyalectaceae AF465440 AY300880
Cameronia pertusarioides 6504 - JX977114 JX977110
Cameronia pertusarioides 6505 - JX977115 JX977111
Cameronia pertusarioides 6506 - JX977116 JX977112
Cameronia tecta - JX977117 JX977113
Chapsa phlyctidioides Graphidaceae JX465300 EU675275
Chapsa pulchra Graphidaceae EU075619 EU075571
Coccomycetella richardsonii Odontotremataceae HM244761 HM244737
Coccotrema cucurbitula Coccotremataceae AF274092 AF329161
Coccotrema pocillarium Coccotremataceae AF274093 AF329166
Coenogonium leprieurii Coenogoniaceae AF465442 AY584698
Coenogonium luteum Coenogoniaceae AF279387 AY584699
Coenogonium pineti Coenogoniaceae AY300834 AY300884
Cryptodiscus pallidus Stictidaceae FJ904677 FJ904701
“Cryptodiscus” rhopaloides - FJ904685 FJ904707
Dibaeis baeomyces Icmadophilaceae AY789291 AY584704
Diploschistes cinereocaesius Graphidaceae AY300835 AY300885
Diploschistes scruposus Graphidaceae AF279389 AY584692
Dyplolabia afzelii Graphidaceae HQ639628 HQ639594
Elixia flexella - AY853368 AY853320
Fissurina insidiosa Graphidaceae DQ973045 DQ972995
Glyphis cicatricosa Graphidaceae HQ639630 HQ639610
Graphis scripta Graphidaceae AY853322 AY853370
Gregorella humida Arctomiaceae AY853329 AY853378
Gyalecta flotowii Gyalectaceae AY300838 AY300889
Gyalecta hypoleuca Gyalectaceae AF465453 HQ659180
Gyalecta truncigena Gyalectaceae HM244766 HM244743
Gyalecta ulmi Gyalectaceae AF465463 AY300888
Gyalectaria gyalectoides Coccotremataceae GU980983 GU980975
Gyalectaria jamesii Coccotremataceae GU980984 GU980976
“Gyalidea”praetermissa - HM244768 HM244745
Hymenelia lacustris Hymeneliaceae AY853371 AY853323
Icmadophila ericetorum Icmadophilaceae DQ883694 DQ986897
Lobothallia radiosa Megasporaceae DQ780306 DQ780274
Myriotrema olivaceum Graphidaceae EU075627 EU075579
Nadvornikia hawaiiensis Graphidaceae AY605080 EU075581
Ocellularia chiriquiensis Graphidaceae EU075629 EU075582
Ocellularia endoxantha Graphidaceae AY605082 EU075589
Ochrolechia androgyna Ochrolechia AY300846 AY300897
Ochrolechia balcanica Ochrolechia AF329171 AF329170
Ochrolechia frigida Ochrolechia AY300847 AY300898
Ochrolechia oregonensis Ochrolechia DQ780308 DQ780276
Ochrolechia pallescens Ochrolechia DQ780310 DQ780277
Ochrolechia parella Ochrolechia AF274097 AF320173
Ochrolechia peruensis Ochrolechia DQ780311 DQ780279
Ochrolechia turneri Ochrolechia AY568002 AY567982
Ochrolechia yasudae Ochrolechia DQ986776 DQ986902
Ochrolechia sp. Ochrolechia DQ986777 DQ986886
Odontotrema phacidiellum Odontotremataceae HM244769 HM244748
Odontotrema sp. Odontotremataceae HM244772 HM244751
Orceolina antarctica Trapeliaceae AY212852 AF274115
Orceolina kerguelensis Trapeliaceae AY212830 AF381561
Paschelkiella pini Stictidaceae HM244762 HM244738
“Pertusaria” albescens Variolaria-group AF329176 AF329175
“Pertusaria” amara Variolaria-group AF274101 AY300900
Pertusaria coccodes Pertusariaceae AF2741095 AY567984
“Pertusaria”corallina Variolaria-group AY300850 AY300901
“Pertusaria” corallophora Variolaria-group DQ780316 DQ780285
Pertusaria coronata Pertusariaceae AY300851 AY300902
Pertusaria gibberosa Pertusariaceae DQ780322 DQ780289
Pertusaria lecanina Pertusariaceae AF274296 AY567991
Pertusaria leioplaca Pertusariaceae AY300852 AY300903
“Pertusaria” mammosa Variolaria-group AY212831 AY212854
Pertusaria mesotropa Pertusariaceae DQ780325 DQ780292
“Pertusaria“ophthalmiza Variolaria-group AY568006 AY567993
Pertusaria paramerae Pertusariaceae DQ780326 DQ780293
Pertusaria pertusa Pertusariaceae AF279300 AF381565
Pertusaria plittiana Pertusariaceae DQ780328 DQ780294
Pertusaria pustulata Pertusariaceae DQ780332 DQ780297
“Pertusaria” scaberula Variolaria-group AF274099 AF431959
“Pertusaria” subventosa Variolaria-group AY300854 AY300905
Phlyctis agelaea Phlyctidaceae AY853381 AY853332
Phlyctis argena Phlyctidaceae DQ986771 DQ986880
Phyllobaeis erythrella - DQ986780 DQ986888
Placopsis cribellans Trapeliaceae DQ871010 DQ871018
Placopsis gelida Trapeliaceae AY212836 AY212859
Placopsis santessonii Trapeliaceae AY212845 AY212867
Placynthiella icmalea Trapeliaceae AY212846 AY212870
Placynthiella uliginosa Trapeliaceae DQ986774 DQ986877
Protothelenella corrosa Protothelenellaceae AY607734 AY607746
Protothelenella sphinctrinoidella Protothelenellaceae AY607735 AY607747
Pycnotrema pynoporellum Graphidaceae JX421615 JX421295
Rhexiophiale rhexoblephara - AY853391 AY853341
Schizoxylon albescens Stictidaceae DQ401144 DQ401142
Siphula ceratites Icmadophilaceae AY853394 AY853344
Schaereria corticola - AY300909 AY300859
Stegobolus subcavatus Graphidaceae EU075641 EU075595
Stictis populorum Stictidaceae AY527327 AY300882
Stictis radiata Stictidaceae AY300864 AY584727
Thamnolia vermicularis Icmadophilaceae AY853345 AY853395
Thecaria quassiicola Graphidaceae HQ639667 JF828971
Thelotrema lepadinum Graphidaceae AY300866 AY300916
Thelotrema subtile Graphidaceae DQ871013 DQ871020
Thelotrema suecicum Graphidaceae AY300867 AY300917
Topeliopsis decorticans Graphidaceae EU075654 EU075609
Trapelia chiodectonoides Trapeliaceae AY212847 AY212873
Trapelia placodioides Trapeliaceae AF274103 AF431962
Trapeliopsis flexuosa Trapeliaceae AF274118 AY212875
Trapeliopsis granulosa Trapeliaceae AF274119 AF381561
Trapeliopsis percrenata Trapeliaceae AF279302 AY212876
Umbilicaria crustulosa Umbilicariaceae AY300869 AY300919
Umbilicaria decussata Umbilicariaceae HM161603 HM161628
Umbilicaria hyperborea Umbilicariaceae AY853399 AY853349
Varicellaria hemisphaerica Varicellaria AF381563 AF381556
Varicellaria lactea Varicellaria AF381557 AF381564
Varicellaria velata Varicellaria AY300855 AY300906
Wawea fruticulosa Arctomiaceae DQ007347 DQ871023
Sequence alignments and phylogenetic analysis

We assembled partial sequences using Geneious Pro 5.4.3 (Drummond et al. 2011) and edited conflicts manually. Alignments were done using Clustal W (Thompson et al. 1994). Ambiguously aligned regions were removed manually. The single locus and concatenated alignments were analyzed by maximum likelihood (ML) and a Bayesian approach (B/MCMC). To test for potential conflict, ML bootstrap analyses were performed on the individual data sets, and 75% bootstrap consensus trees were examined for conflict (Lutzoni et al. 2004). Maximum likelihood analyses were performed using the program GARLI (Zwickl 2006), employing the general time reversible model of nucleotide substitution (Rodriguez et al. 1990), including estimation of invariant sites, and assuming a discrete gamma distribution with six rate categories as in Lumbsch et al. (2007b). Bootstrapping (Felsenstein 1985) was performed based on 2000 replicates. The B/MCMC analysis was conducted on the concatenated data set using MrBayes 3.1.2 (Huelsenbeck and Ronquist 2001), with the same substitution model as in the ML analysis. The dataset was partitioned into two (mtSSU, nuLSU) and each part was allowed to have its own parameters (Nylander et al. 2004). A run with 20, 000, 000 generations, starting with a random tree and employing 4 simultaneous chains, was executed. Every 100th tree was saved. The first 500, 000 generations (i.e. the first 5000 trees) were deleted as the “burn in” of the chain. We used AWTY (Nylander et al. 2007) to compare split frequencies in the different runs and to plot cumulative split frequencies to ensure that equilibrium was reached. Of the remaining trees, a majority rule consensus tree with average branch lengths was calculated using the sumt option of MrBayes. Posterior probabilities were obtained for each clade. Only clades that received bootstrap support equal or above 70% under ML and posterior probabilities ≥ 0.95 were considered as strongly supported. Phylogenetic trees were depicted using the program FigTree 1.3.1 (Rambaut 2009).

Results and discussion

Eight new sequences were generated for this study and aligned with sequences downloaded from Genbank (Table 1). The single gene locus trees did not show any conflicts and hence the concatenated data set was analyzed. Our combined data set included 1313 unambiguously aligned positions, 370 of which were constant. The ML tree had a likelihood value of –26318.540 and in the B/MCMC analysis of the combined data set, the likelihood parameters in the sample had the following mean (Variance): LnL = -27045.138 (0.35). The ML tree and the tree from the B/MCMC tree sampling were almost identical, with no differences in well-supported clades. Furthermore, taxon sampling was very similar to that of previous studies focusing on the phylogeny of Ostropomycetidae (Baloch et al. 2010; Lumbsch et al. 2007a; Lumbsch et al. 2007b; Wedin et al. 2005). Thus, only a simplified ML tree, with samples of well-supported families, genera or generic groups collapsed, is shown here (Fig. 1). Individual OTUs are shown only for the species of Cameronia and its sister groups. In our analysis, the four samples of the two Cameronia species form a strongly supported, monophyletic group within the well-supported Ostropomycetidae, confirming the monophyly of the genus and its placement in Ostropomycetidae. The genus Cameronia is another example of a group of lichenized ascomycetes with perithecioid ascomata in this subclass, with others being Porinaceae (Baloch and Grube 2006; Grube et al. 2004), Protothelenellaceae and Thelenellaceae (Schmitt et al. 2005). There are additional families in this subclass that also include taxa with more or less perithecioid ascomata, such as Coccotremataceae, Gyalectaceae, Pertusariaceae and Graphidaceae (Baloch et al. 2010; Lumbsch and Schmitt 2002; Lumbsch et al. 2001; Rivas-Plata et al. 2012; Rivas-Plata and Lumbsch 2011; Schmitt et al. 2010; Schmitt and Lumbsch 2004). The diversity of ascomatal morphologies in this subclass has been linked to the hemiangiocarpous type of ascoma development in the group as a whole (Schmitt et al. 2009).

Figure 1.

Phylogenetic placement of Cameronia as inferred from a concatenated alignment of mtSSU and nuLSU DNA sequences. This is a simplified cartoon of the optimal tree under maximum likelihood with well supported families and species groups collapsed that were shown in previous studies (Baloch et al. 2010; Lumbsch et al. 2007a; Lumbsch et al. 2007b; Wedin et al. 2005). Asterisks indicate branches with likelihood bootstrap support values above 70% and posterior probabilities equal or above 0.95.

The backbone of the Ostropomycetidae tree largely lacks support and the relationships of Cameronia within Ostropomycetidae remain unclear. Cameronia is the sister-group of Baeomycetaceae (Ainoa, Baeomyces, Phyllobaeis) but this relationship lacks support. This clade forms a sister-group to a well-supported clade that includes Anzina and Protothelenellaceae, but again, this relationship lacks support.

Although the molecular data support the placement of Cameronia in Ostropomycetidae, they fail to identify any close relatives of the genus, which is also reflected in the similarities of Blast searches of the newly generated sequences (maximal identity - nuLSU: 94%, mtSSU: 93%). Cameronia is distinguished by several characters that are generally used to characterize families, as shown in Table 2 where salient features of Cameronia and other families of Ostropomycetidae with perithecioid ascomata (Porinaceae, Protothelenellaceae, Thelenellaceae) are compared. The ascus type is very different from any of the other perithecioid Ostropomycetidae and also different from the apotheciate Baeomycetaceae, which have cylindrical asci (Gierl and Kalb 1993). Nor is the rudimentary exciple seen in Cameronia found in any of the other perithecioid families. Morphologically, the most similar family in Ostropomycetidae is Protothelenellaceae, with which Cameronia shares a hamathecium of richly branched paraphysoids and a lack of periphyses. However, Prothelenellaceae have a different exciple, different asci with an amyloid apical apparatus in the tholus and an ocular chamber, and halonate ascospores. Furthermore, Protothelenellaceae form a well-supported clade with Anzina (Fig. 1) and are only distantly related to Cameronia. The isolated position of Cameronia is consistent with the hypothesis that this genus is a case of paleoendemism. It will be an exciting project to test this hypothesis at a later stage when more sequence data from Ostropomycetidae become available.

Table 2.

Diagnostic features of families with perithecioid ascomata in Ostropomycetidae (Baloch and Grube 2006; Fryday and Coppins 2004; Grube et al. 2004; Kantvilas 2012; Mayrhofer 1987b, 2002; Mayrhofer and Poelt 1985; McCarthy 1995; McCarthy 2000).

Characters Cameronia Porinaceae Protothelenellaceae Thelenellaceae
Proper exciple rudimentary Well developed, consisting of periplectenchymatous cells Well developed, consisting of periplectenchymatous to isodiametric cells Well developed, consisting of periplectenchymatous cells
Hamathecium Richly branched, anastomosing paraphysoids, no periphyses Simple to sparsely branched Paraphyses, no periphyses Richly branched, anastomosing paraphysoids, no periphyses Richly branched, anastomosing paraphysoids, periphyses present
Asci Broadly obovate cylindrical cylindrical cylindrical
Tholus Well-developed Poorly developed Well-developed Poorly developed
Ascus amyloidity Outer wall hemiamyloid, tholus non-amyloid Non-amyloid Outer and wall and tholus amyloid Non-amyloid
Ocular chamber - - + +/-
Ascospores Hyaline, non-halonate, thick-walled, muriform Hyaline, halonate, thin- to thick-walled, transversely septate to muriform Hyaline, halonate, thick-walled muriform Hyaline to brownish, halonate, thin-walled, muriform
Chemistry Dibenzofuranes, triphenyl Nil or pigments nil nil

Given the dissimilarity in morphological characters and the lack of close relatives in the phylogenetic tree, we propose a new family Cameroniaceae below to accommodate the genus Cameronia. The new family is placed in Ostropomycetidae with unclear ordinal position.

Cameroniaceae Kantvilas & Lumbsch, fam. nov.

Mycobank no. MB802404


Cameronia Kantvilas, Lichenologist 44: 92. 2012.


Thallus crustose, photobiont a coccoid green alga. Ascomata perithecioid, immersed in the thallus, proper exciple rudimentary, hamathecium consisting of richly branched, anastomosing paraphysoids, inspersed with oil droplets, containing hymenial algae, periphyses absent. Asci broadly obovate, with outer wall hemiamyloid and with a well-developed, non-amyloid tholus; ocular chamber lacking. Ascospores hyaline, non-halonate, eumuriform. Conidiomata immersed in the thallus, forming baciliform to bone-shaped conidia.


This study was supported by the NSF grant “ATM – Assembling a taxonomic monograph: The lichen family Graphidaceae” (DEB-1025861). The laboratory work was done at the Pritzker Laboratory for Molecular Systematics at the Field Museum. For companionship in the field in quest of fresh material for analysis, GK thanks Brigitte de Villiers.

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