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Systematics |
2Departament de Biologia Vegetal (Unitat de Botànica), Facultat de Biologia, Universitat de Barcelona, Av. Diagonal 645, 08028 Barcelona, Spain; 3Department of Biology, Duke University, Durham, North Carolina 27708-0338 USA
Received for publication October 29, 2002. Accepted for publication February 14, 2003.
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ABSTRACT |
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 TOP ABSTRACT INTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Key Words: ambiguous alignments • Caloplaca • Fulgensia • internal transcribed spacer • lichen-forming fungi • Markov chain Monte Carlo Bayesian inference • molecular phylogenetics • Teloschistaceae • Xanthoria
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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.
The genus was described by Massalongo and de Notaris (Massalongo, 1855
) based on Fulgensia fulgens (Sw.) Elenkin, i.e., mainly comprising taxa with non-polarilocular, mostly simple ascospores, and with more or less pruinose and yellowish thalli containing anthraquinones. Zahlbruckner (1926
, 1931
) did not recognize Fulgensia as a distinct genus and subsumed it within Caloplaca Th. Fr. as a separate section. Dodge and Baker (1938)
, Räsänen (1943)
, and Rudolph (1955)
revived the use of the name Fulgensia at the genus level.
Poelt (1965)
redefined the previous delimitation of Fulgensia by including, in addition to taxa with simple ascospores, taxa with typical one-septate non-polarilocular ascospores. Fulgensia sensu Poelt contained 10 species distributed into two subgenera: Fulgensia subgen. Fulgensia Poelt and subgen. Candelariopsis (Sambo) Poelt. The former included six species: F. delphinensis Poelt, F. desertorum (Tomin) Poelt, F. bracteata (Hoffm.) Räsänen, F. subbracteata (Nyl.) Poelt, F. fulgens (Sw.) Elenkin, and F. fulgida (Nyl.) Szatala. This subgenus was characterized mainly by simple or one-septate ascospores, a thallus with a false upper cortical layer, and by their attachment to the substratum with a felt of hyphae (Poelt, 1965
). The subgenus Candelariopsis included four taxa: F. australis (Arnold) Poelt, F. chanousiae (Sambo) Poelt, F. schistidii (Anzi) Poelt, and F. pruinosa (Körb.) Poelt. These species have one-septate ascospores, a thallus with a true upper cortical layer, and are attached to the substratum by hyphal strands or by their lower cortex (Poelt, 1965
). Subsequently, three new Fulgensia species were described: F. klementii Kalb (Kalb, 1970
), F. poeltii Llimona (Llimona, 1974
), and F. canariensis Follmann (Follmann and Poelt, 1981
).
Gilbert (1978)
applied Poelt's concepts to Fulgensia populations found in the British Isles. He asserted the complexity of this genus by remarking that most of the species exhibit a large variability, implying the existence of several potential infraspecific taxa. Poelt's circumscription of Fulgensia has not always been followed. Some contemporary authors (Nowak and Tobolewski, 1975
; Oksner, 1993
) still recognize members of the Candelariopsis group as being Caloplaca species.
Apart from Poelt (1965)
, no further revision of the group had been made until the work of Westberg and Kärnefelt (1998)
, who recognized four groups within Fulgensia: group A (F. australis and F. chanousiae), group B (F. canariensis), group C (F. schistidii) and group D (F. bracteata, F. delphinensis, F. desertorum, F. fulgens, F. fulgida, F. klementii, F. poeltii, F. pruinosa, and F. subbracteata). In their study of Fulgensia and Caloplaca, Poelt (1965)
and Kärnefelt (1989)
came to the conclusion that species from the Caloplaca aurea group (not sampled here) were most similar to Fulgensia (Candelariopsis), the main differential feature being the type of ascospore septation. Based on the structure of the upper thallus cortex, Kärnefelt (1989)
and Westberg and Kärnefelt (1998)
proposed close affinities between species from the C. aurea group (e.g., C. paulii Poelt) and F. canariensis and F. schistidii. However, Navarro-Rosinés et al. (2000)
suggested that C. aurea (Schaer.) Zahlbr. was more similar to subgenus Fulgensia than to subgenus Candelariopsis. Furthermore, the recently described C. cancarixiticola Nav.-Ros., Egea & Llimona shares several morphological features with F. australis, F. canariensis, and F. schistidii (Navarro-Rosinés et al., 2000
).
In spite of all these morphological studies and the evident connections between Fulgensia and Caloplaca, only one molecular phylogenetic study of Fulgensia has been carried out (Kasalicky et al., 2000
). Unfortunately, only two Caloplaca species (C. flavorubescens [Huds.] Laundon and C. cerina [Hedw.] Th. Fr.) and one Xanthoria (Fr.) Th. Fr. species (X. parietina [L.] Th. Fr.) were included to represent taxa outside Fulgensia. This sampling was sufficient to demonstrate the polyphyletic nature of Fulgensia, but was insufficient to reveal relationships of its separate lineages within the broad context of the Teloschistaceae.
During the molecular and morphological revision of the lobate Caloplaca (Gasparrinia sect.) by the first author, the need to clarify the delimitation between Caloplaca and Fulgensia became necessary. We report here the results of a phylogenetic study assessing relationships of Fulgensia species within the broader context of the Teloschistaceae. The conclusions presented here are based on sequences from the internal transcribed spacer (ITS) region for 58 specimens, representing 33 species (36 taxa) belonging to three genera (Caloplaca, Fulgensia, and Xanthoria) within the Teloschistaceae.
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MATERIALS AND METHODS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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; Kasalicky et al., 2000
; Lohtander et al., 2000
; Martín and Winka, 2000
; Dyer and Murtagh, 2001
; Kroken and Taylor, 2001
). Eight new ITS sequences were generated for this study. After comparing ITS sequences from a number of species belonging to different families with our alignment of ingroup sequences, four outgroup species were selected: Protoparmelia badia (Hoffm.) Haf. (Lecanoraceae), Letharia vulpina (L.) Hue, L. columbiana (Nutt.) J. W. Thomson, and Usnea arizonica Mot. (Parmeliaceae). See Supplemental Data accompanying the online version of this paper for voucher information and GenBank accession numbers for the 62 ITS sequences included in this study.
DNA isolation and sequencing
Genomic DNA was obtained from fresh samples and herbarium specimens (the oldest specimen was collected in 1995). DNA was isolated using the Puregene Kit (GENTRA Systems, Minneapolis, Minnesota, USA) following the manufacturer's protocol for filamentous fungi. DNA concentration was determined by visual comparison with positive control (
100 ladder, concentration 10, 20, 40 ng) on an ethidium-bromide-stained agarose gel. Symmetric polymerase chain reactions (PCR) were prepared for a 50.0 µL final volume containing 31.7 µL of sterile double-distilled water, 5.0 µL of 10x Taq polymerase reaction buffer (Boehringer-Mannheim, Indianapolis, Indiana, USA), 5.0 µL of 2.5 mmol/L dNTPs, 0.3 µL of Taq DNA polymerase (Boehringer-Mannheim), 2.5 µL for each of the 10 µmol/L primers ITS1F or ITS5 and ITS4 (Gardes and Bruns, 1993
; White et al., 1990
), 1.5 µL of 10 mg/mL bovine serum albumin (BSA; BioLabs, Beverly, Massachusetts, USA), 0.5 µL of 50 mmol/L MgCl2, and 1 µL of template genomic DNA. The PCR was performed on Peltier Thermal Cyclers PTC-200 (MJ Research, Boston, Massachusetts, USA) under the following conditions: one cycle of 1 min at 95°C linked to 40 cycles of 1 min at 95°C, 45 s at 52°C, and 2 min at 72°C, with the last step increased by increments of 5 s for the last 15 cycles. A final extension step of 10 min at 72°C was added, after which the samples were kept at 4°C. The PCR products were purified using GELase Agarose Gel-Digesting Preparation (Epicentre Technologies, Madison, Wisconsin, USA) following the manufacturer's instructions or low-binding regenerated cellulose 30 000 nominal molecular weight limit (NMWL) filter units (Millipore, Bedford, Massachusetts, USA). Both strands of the purified PCR products were sequenced using PCR primers used for the symmetric amplification and primers 5.8S and 5.8SR (Vilgalys and Hester, 1990
). Sequencing reactions were prepared in 10 µL final volume using BigDye Terminator (ABI PRISM, Perkin-Elmer Biosystems, Wellesley, Massachusetts, USA) and following the manufacturer's instructions. Sequenced products were precipitated with 10 µL of deionized sterile water, 2 µL of 3 mol/L sodium acetate, and 50 µL of 95% ethanol. Polyacrylamide gel electrophoresis was conducted using Long Ranger Singel packs (FMC BioProducts, Rockland, Maryland, USA) and an ABI 377 automated DNA sequencer (Perkin-Elmer, Applied Biosystems, Wellesley, Massachusetts, USA). Sequence fragments were subjected to BLAST searches to verify their identity and assembled using Sequencher version 3.0 (Gene Codes Corporation, Ann Arbor, Michigan, USA).
Sequence alignment
Sequences were aligned using Sequencher 3.0. The manual alignment was carefully inspected for the presence of ambiguously aligned regions caused by the insertion of gaps. The alignment was done separately in two blocks because of the difficulty of aligning two different groups of sequences. Xanthoria fallax (Hepp) Arnold was the only species alignable with both groups and was used to facilitate the final and global manual alignment. The delimitation of ambiguous regions, their unequivocal coding, and the elaboration of symmetric step matrices for each of these coded characters were generated using the program INAASE 2.3b (Lutzoni et al., 2000
). The alignment is available in a Nexus format upon request to E.G. or F.L.
Phylogenetic analyses
All phylogenetic searches were performed with PAUP* 4.0b8 (Swofford, 2001
) using maximum parsimony (MP) and maximum likelihood (ML) as optimization criteria. The same program was used for all bootstrap analyses. Constant sites and ambiguously aligned sites were removed from all analyses.
Maximum-parsimony (MP) analyses
Three different MP searches (MP1–MP3) were carried out.
A first MP analysis (MP1) was executed using exclusively unambiguously aligned sites. Changes among character states (including gaps as a fifth character state) were subjected to symmetric step matrices constructed as follows. The options Show character status/full details/hide excluded characters options from the Data menu in PAUP* were implemented. From the resulting table, the column States showing all nucleotide states found at each of the unambiguously aligned and nonconstant sites was saved as a separate text file. This file was used as an input file for the program STMatrix 2.1 (written by S. Zoller and available upon request from S.Z. or F.L.), which generates a step matrix (in Nexus format) by calculating frequencies of reciprocal changes from one state to another and converting them into costs of changes using the negative natural logarithm of the frequencies (Felsenstein, 1981
; Wheeler, 1990
).
The second MP search (MP2) was restricted to unequivocally coded (INAASE) characters replacing ambiguously aligned regions. Only coded characters with no more than 32 character states were subjected to the MP2 search. A total of eight coded characters out of 16 (corresponding to the 16 ambiguously aligned regions we delimited in this ITS alignment) fit this criterion, forming eight new (INAASE) characters. All pair-wise transformation costs were equal for one of these eight characters. The seven other coded characters (with consistent pair-wise sequence comparisons) were each subjected to a specific step matrix taking into account the optimal number of steps to transform one ambiguous sequence into another.
Before combining the unambiguously aligned sites with the eight coded (INAASE) characters for the MP3 analysis, global phylogenetic congruence was assessed for these two data partitions. This was done by inspecting internodes with bootstrap scores above 70% resulting from the separate MP1 and MP2 analyses (Mason-Cramer and Kellogg, 1996
). Conflicts between the two data partitions would be considered significant only if the two bootstrap analyses provided support 
70% for two different phylogenetic relationships for the same set of taxa (one being monophyletic and the other being nonmonophyletic).
All three MP searches were performed using heuristic searches with 1000 random-addition-sequences (RAS), tree bisection-reconnection (TBR) branch swapping, Multrees option in effect, and collapsing branches with maximum branch length equal to zero. The MP1 and MP2 searches were done in two steps. The goal of the first step was to find the length of the most parsimonious trees. This was done by saving only one tree for each RAS. The goal of the second step was to find all equally most parsimonious trees. This was done by saving all trees only when swapping on a tree equal or shorter than the shortest tree found in the first step. The MP3 was conducted in one step by saving all trees as soon as TBR swapping was initiated. Internode confidence was assessed by bootstrap analyses (Felsenstein, 1985
) with full heuristic searches, 10 000 parsimony bootstrap replicates, using two RAS per bootstrap replicate and by saving no more than 10 trees per bootstrap replicate.
Maximum-likelihood (ML) analyses
Selection of the nucleotide substitution model and the estimation of most parameters were done with hierarchical likelihood ratio testing (HLRTs) using the program Modeltest 3.04 PPC (Posada and Crandall, 1998
). The ML search was performed using the HKY substitution model (Hasegawa-Kishino-Yano, 1985
) with base frequencies (A = 0.1539, C = 0.2704, G = 0.2263, T = 0.3494), ti/tv ratio = 3.2956, and unequal rates among sites. Rates for variable sites assumed to follow a gamma distribution with shape parameter = 1.9549 for an optimal number of rate categories equal to 3. The ML heuristic search was conducted on the same data set subjected to the MP1 search, with 1000 RAS, number of rearrangements limited to 20 000/RAS, TBR branch swapping, Multrees option in effect, and collapsing branches with maximum branch length equal to zero. The ML analysis was divided in three separate searches (300, 300, and 400 RAS, respectively).
We used a Bayesian Markov chain Monte Carlo (B/MCMC) method (Larget and Simon, 1999
) to assess the level of confidence for relationships revealed by our ML search. The same model and number of rate categories as used in the ML search was used for the B/MCMC analyses. The tree sampling using this approach was done with MrBayes 1.11 (Huelsenbeck, 2000
). The MCMC algorithm ran for 2 000 000 generations with four incrementally heated chains, starting from random trees and sampling one out of every 100th generation with DNA substitution parameters updated during the search. A majority-rule consensus tree was calculated with PAUP* from the last 15 000 out of the 20 000 trees sampled. The first 5000 trees (burnin) were excluded to avoid trees that might have been sampled prior to convergence of the Markov chains.
We repeated the B/MCMC tree sampling as described to confirm the result from the first Bayesian analysis. The final majority-rule consensus tree was obtained by pooling all trees selected from both runs (30 000 trees). The posterior probability of each topological bipartition was estimated by the frequency of these bipartitions across all 30 000 trees sampled. Internodes with posterior probabilities 95% were considered statistically significant.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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The unequally weighted MP1 search revealed 575 equally most parsimonious trees of 288.19 steps (Fig. 1). The unequally weighted MP2 search revealed 5712 equally most parsimonious trees in two islands. One island of 4032 trees was hit 101 times and the second island of 1680 trees was hit 49 times. The score of the best tree was 173.00 steps (Fig. 2).
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50% generated by the MP2 bootstrap analysis, 11 internodes had bootstrap values increased by an increment of at least 10% when compared to the MP1 bootstrap values, and six of these bootstrap values went from <70% to 70% (Fig. 2). Although topologies obtained from the separate analyses of unambiguous sites and INAASE characters were considerably different, no conflict was detected using the reciprocal 70% bootstrap criterion (see Materials and Methods), and, therefore, the two data sets were combined.
A total of 12 equally most parsimonious trees was found in one island that was hit 1000 times with the MP3 search based on combined unambiguously aligned sites and INAASE characters (Fig. 3). The score of the best tree was 473.70 steps. The inclusion of the eight ambiguously aligned regions as coded characters explains the increase in resolution from 575 to 12 equally most parsimonious trees and the faster computing time when the analysis was performed on the combined data set. The number of internodes with bootstrap support 70% went from 13, when the analysis was restricted to unambiguously aligned sites (MP1), to 25 when the eight INAASE characters were added to these unambiguous sites (Figs. 1 and 3).
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94%) with B/MCMC (Fig. 4) compared to 25 (bootstrap proportion [BP] 70%) for MP3 (Fig. 3). This extra level of support comes from the phylogenetic signal that was recovered with the eight INAASE characters from ambiguously aligned portions of the alignment (Fig. 2). When the MP bootstrap analysis was performed on the same data set that was used for the B/MCMC analysis, i.e., without the eight INAASE characters, internodes with high support were virtually the same (compare Fig. 1 with Fig. 4). The B/MCMC approach did provide significant support for two internodes that never received bootstrap support 70% (Fig. 4).
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, our rooting of the ingroup is in total agreement with the result shown in their Fig. 4.
Phylogenetic relationships
Our results suggest that Caloplaca, Xanthoria, and Fulgensia are part of two main lineages within the Theloschistaceae: lineage 1 (PP = 99%, Fig. 4) and lineage 2 (BP = 89%, Fig. 3). Because Xanthoria and most lobate Caloplaca species are restricted to lineage 1, this lineage seems to include fewer species and to be more homogeneous than lineage 2. By including most members of Fulgensia and several representatives of at least five groups of Caloplaca (sensu Clauzade and Roux, 1985
), lineage 2 is the most phenotypically diverse.
Fulgensia has three independent origins, confirming the polyphyly of this genus by Kasalicky et al. (2000)
. One of these origins gave rise to the largest group of Fulgensia species (BP = 95%, Fig. 3; PP = 98%, Fig. 4) and corresponds to the Fulgensia sensu stricto (s.s.) group proposed by Kasalicky et al. (2000)
. Fulgensia s.s. and F. australis are part of lineage 2 of the Teloschistaceae, whereas F. schistidii is part of lineage 1 of this family.
Our B/MCMC analysis shows that Fulgensia s.s. is nested within a subgroup of Caloplaca part of lineage 2. Exact relationships among the main species groups within lineage 2 remain uncertain with ITS. However, a closer (monophyletic) relationship of Fulgensia s.s. to Caloplaca gloriae Llimona & Werner, C. flavovirescens (Wulfen) DT. & Sarnth, and C. flavorubescens was found to be statistically significant (PP = 98%, Fig. 4).
The phylogenetic position of Fulgensia australis within lineage 2 is also uncertain. Both the MP and the ML analyses suggest a close relationship with C. dolomiticola (Hue) Zahlbr. and C. alpigena Poelt ined. (current name: C. macrocarpa (Anzi) Zahlbr.), but virtually without any support (Figs. 3 and 4). Caloplaca variabilis (Pers.) Müll. Arg. was always at the base of lineage 2 in the combined analyses, but did not receive statistical support.
In general, the level of uncertainty is also high for relationships among main groups within lineage 1, except for the Xanthoria fallax group, which was constantly found to be part of the first divergence that took place within this lineage (PP = 94%, Fig. 4; BP = 72%, Fig. 3). This result is in agreement with the ITS tree from Arup and Grube (1999)
.
Fulgensia schistidii is most likely to form a monophyletic group with members of the Caloplaca saxicola group (BP = 73%, Fig. 3), rather than being nested within Fulgensia where it has been classified traditionally (Poelt, 1965
) or being sister to Xanthoria parietina according to Kasalicky et al. (2000)
. In agreement with Arup and Grube (1999)
, C. marina (Wedd.) Zahlbr. and C. maritima B. de Lesd. are sister species with strong support. However, by adding Fulgensia species (Kasalicky et al., 2000
) to the data set of Arup and Grube (1999)
, we lost the high support for the sister relationship of C. lithophila H. Magn. to the C. marina group.
Our study revealed a sister relationship between X. sorediata (Vain.) Poelt and X. elegans (Link) Th. Fr. (BP = 71%, Fig. 3), as well as between X. parietina and X. resendei Poelt & Tav. (BP = 79%, Fig. 3). However, one specimen of "X. elegans" (individual 1) was always found to be sister to C. arnoldii (BP = 63%, Fig. 3; PP = 95%, Fig. 4). This specimen of "X. elegans" with "compact thalli" was collected in eastern Antarctica and was part of a phylogenetic study by Dyer and Murtagh (2001)
. Based on their Figs. 2 and 4, this "Xanthoria elegans" with compact thalli is very likely a misidentified member of the C. saxicola group (GenBank AF278753). Even if we reassign "Xanthoria elegans 1" to Caloplaca saxicola sensu lato, our results clearly show that Xanthoria and Caloplaca do not form monophyletic groups (Figs. 3 and 4).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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Adequacy of ITS in resolving relationships within the Teloschistaceae
One important finding to emerge from this study is that in spite of the small number (eight) of ambiguous regions that we were able to convert to characters with INAASE, these eight characters alone were able to generate a high level of resolution. However, the highest level of confidence with MP was obtained when these eight INAASE characters were added to the data matrix containing the nonambiguously aligned sites. Doing so, the number of internodes with bootstrap support 70% was more than doubled compared to the MP1 analysis.
It is clear that large ITS data sets within the Teloschistaceae will greatly benefit from methods like INAASE that are designed to recover phylogenetic signal from these ambiguously aligned regions. In this particular case, using B/MCMC, which seems to be a more statistically powerful method to assess the level of phylogenetic uncertainty (Alfaro et al., 2002
; Kauff and Lutzoni, 2002
), is not superior to using parsimony bootstrap with INAASE characters. This is likely to be true for alignments with high percentages of sites that are ambiguously aligned, until B/MCMC methods and ML searches can accommodate multiple models simultaneously, including models for characters with character states other than the usual four nucleotides (e.g., INAASE or morphological characters). It is possible that ITS alone could provide sufficient phylogenetic information to fully resolve relationships within the Teloschistaceae and provide high support values for most internodes, if a new method could capture phylogenetic signal from all ambiguously aligned regions instead of being restricted to eight of the 16 regions we delimited for this study.
Phylogenetic relationships of Fulgensia species within the Teloschistaceae
Fulgensia pruinosa had been included in subgenus Candelariopsis with F. australis, F. chanousiae, and F. schistidii by Poelt (1965)
. Westberg and Kärnefelt (1998)
, Kasalicky et al. (2000)
, and this study demonstrate that F. pruinosa belongs to the subgenus Fulgensia, corresponding to Fulgensia s.s. This group is characterized by a homogeneous cortical layer and abundant pruina.
As for Kasalicky et al. (2000)
, our analyses could not distinguish "F. desertorum" from F. bracteata. The high similarity of the ITS and LSU sequences suggest that more characters are needed to resolve this relationship. We believe that because of the overall phenotypic similarity of "F. desertorum" to F. fulgens, this odd result may also be explained by misidentification of the two specimens used by Kasalicky et al. to represent "F. desertorum." Typical F. desertorum is xerophilous and of irano-turanian distribution (X. Llimona, personal communication), whereas specimens used by Kasalicky et al. (2000)
have a very different ecology and were collected in Norway and northern Italy.
We confirm the conclusion by Westberg and Kärnefelt (1998)
and Kasalicky et al. (2000)
that Fulgensia australis does not belong to Fulgensia s.s. However, its phylogenetic placement is still unknown. This might be resolved by including more related taxa. For example, species from the Caloplaca aurea group share several morphological similarities with F. australis, i.e., lack of pruina, presence of septate spores, and saxicolous habit. Additional molecular characters might also be necessary to resolve the phylogenetic relationship of this problematic species.
Because of its placement in lineage 1, F. schistidii appears to be the most distantly related species to Fulgensia s.s. Westberg and Kärnefelt (1998)
already pointed out the particularity of this species, creating a monospecific group for it. Our study revealed a tight relationship between F. schistidii and species of the C. saxicola group. The yellow-orange color, the septate spores, the compact cortex and medulla, as well as the presence of lobes (though sometimes highly reduced) corroborate in part this relationship.
Taxonomic conclusions
The type species of Caloplaca (C. cerina) is nested within lineage 2, together with Fulgensia s.s. and several morphologically diverse Caloplaca groups. Hence, lineage 2 forms a heterogenous group for which it is not yet possible to determine the taxonomic rank without a more extensive taxon sampling and more characters. However, we can conclude that this taxonomic unit contains both Caloplaca s.s. and Fulgensia s.s. We can also conclude that Fulgensia australis (Arnold) Poelt should revert to Caloplaca australis (Arnold) Zahlbr.
Lineage 1 is mostly formed by yellowish-orange lobed species that seem to be centered around subgenus Gasparrinia (Torn.) Th. Fr. (sensu Clauzade and Roux, 1985
). Some of the most extreme forms in this lineage are represented by Xanthoria species, Fulgensia schistidii, and Caloplaca pyracea (Ach.) Th. Fr. Therefore, lineage 1 forms a fairly homogeneous group, especially when compared to lineage 2. In this context, the generic rank could be interpreted as being the most appropriate for lineage 1. Given that Xanthoria parietina, the type of Xanthoria, is part of this lineage, the name Xanthoria is available for lineage 1. This would mean that all Caloplaca species in this lineage and Fulgensia schistidii would have to be transferred to Xanthoria.
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FOOTNOTES |
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5 Author for reprint requests (gaya{at}porthos.bio.ub.es
)
4 Present address: North Carolina Supercomputing Center, PO Box 12889, Research Triangle Park, North Carolina 27709 USA
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LITERATURE CITED |
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TOPABSTRACTINTRODUCTIONMATERIALS AND METHODSRESULTSDISCUSSIONLITERATURE CITED |
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