Research Article
Research Article
New species and records of Chapsa (Graphidaceae) in China
expand article infoMing-Zhu Dou, Min Li, Ze-Feng Jia
‡ Liaocheng University, Liaocheng, China
Open Access


We studied the genus Chapsa in China based on morphological characteristics, chemical traits and molecular phylogenetic analysis. One species new to science (C. murioelongata M.Z. Dou & M. Li) and two records new to China were found (C. wolseleyana Weerakoon, Lumbsch & Lücking and C. niveocarpa Mangold). Chapsa murioelongata sp. nov. is characterised by its lobed thalline margin, orange discs with white pruina, clear hymenium, and submuriform and long ascospores. Chapsa wolseleyana was recombined into Astrochapsa based on phenotypic traits. Sequences of this species are for the first time reported here and phylogenetic analyses of three loci (mtSSU, ITS and nuLSU) supported the position of this species within Chapsa. A key for the Chapsa species known in China is provided.


Ascomycota, lichenized fungi, phylogeny, taxonomy


The lichen genus Chapsa (Graphidaceae) was first established by Massalongo (1860) with C. indica as the type species. This genus was ignored for a long time until 2006, when Frisch re-established Chapsa, based on the Chroodiscus-type apothecia, presence of periphysoids and Chapsa-type paraphyses. Frisch (2006) also provided a detailed description and delimitation of the genus Chapsa, which was widely recognised by subsequent researchers (Mangold 2008; Frisch and Kalb 2009; Rivas Plata et al. 2011; Sipman et al. 2012; Xu et al. 2016). The genus Chapsa was considered to be monophyletic in the beginning (Frisch 2006) but with further research, it was suspected to be polyphyletic (Mangold 2008; Papong et al. 2010). Subsequently, seven genera, Astrochapsa Parnmen, Lücking & Lumbsch, Crutarndina Parnmen, Lücking & Lumbsch, Gintarasia Kraichak, Lücking & Lumbsch, Pseudochapsa Parnmen, Lücking & Lumbsch, Pseudotopeliopsis Parnmen, Lücking & Lumbsch, Myriochapsa M. Cáceres, Lücking & Lumbsch and Nitidochapsa Parnmen, Lücking & Lumbsch were separated from Chapsa, based on a combination of molecular evidence, phenotypic and chemical characteristics (Parnmen et al. 2012, 2013; Kraichak et al. 2013).

Although China is rich in lichenised fungal species (Wei 2020), there are few studies and reports on the genus Chapsa. More than 60 species of Chapsa have been reported in the world, of which only three, C. indica A. Massal, C. mirabilis A. (Zahlbr.) Lücking and C. leprocarpa (Nyl.) Frisch, have so far been found in China (Rivas Plata et al. 2010; Xu et al. 2016; Jia and Lücking 2017; Kalb and Kalb 2017; Wijayawardene et al. 2017; de Lima et al. 2019).

During the study of Chapsa A. Massal. in southern China, one species, C. murioelongata was found new to science, and two species, C. niveocarpa Mangold and C. wolseleyana Weerakoon, Lumbsch & Lücking were found new to China. In our study, 26 sequences were newly generated from freshly collected specimens.

Materials and methods

Morphological and chemical analyses

The specimens were collected from southern China and deposited in the Fungarium, College of Life Sciences, Liaocheng University, China (LCUF). Morphological and anatomical characters of thalli and apothecia were examined and photographed under an Olympus SZX16 dissecting microscope and an Olympus BX53 compound microscope. The lichen secondary metabolites were detected and identified by thin-layer chromatography using solvent C (Orange et al. 2010; Jia and Wei 2016).

DNA extraction, PCR sequencing and phylogenetic analysis

Genomic DNA was extracted from ascomata using the Hi-DNA-secure Plant Kit (Tiangen, Beijing, China) according to the manufacturer’s protocol. The nuLSU, ITS and mtSSU regions were amplified using the primer pair AL2R/LR6 (Mangold 2008, Vilgalys and Hester 1990), ITS1F/ITS4 (Gardes and Bruns 1993, White et al. 1990) and mrSSU1/mrSSU3R (Zoller et al. 1999), respectively. The PCR amplification progress followed Dou et al. (2018) and the PCR products were sequenced by Biosune Inc. (Shanghai). The newly generated sequences were submitted to GenBank (Table 1).

Table 1.

Information for the sequences used in this study. Newly generated sequences are shown in bold.

Species Specimen No. Locality ITS nuLSU mtSSU
Pseudochapsa phlyctidioides Lumbsch 20500d Fiji JX465301 JX421005
Pseudochapsa dilatata Luecking 32101 Venezuela JX421446 JX420981
Pseudochapsa esslingeri Caceres s.n. Brazil JX420983
Pseudochapsa esslingeri Caceres 6006a Brazil JX420984
Pseudochapsa esslingeri Rivas Plata 107C (F) Peru JX420985
Pseudochapsa esslingeri Rivas Plata 809a (F) Peru JX420986
Chapsa alborosella Luecking 31238a Brazil JX421439 JX420972
Chapsa alborosella Luecking 25587 Guatemala JX421440 JX420973
Chapsa soredicarpa Luecking 31200 Brazil JX421462 JX421011
Chapsa soredicarpa Luecking 31240 Brazil JX421463 JX421012
Chapsa sublilacina Luecking RLD056 Mexico HQ639624 HQ639600
Chapsa thallotrema Lucking 32019 Venezuela JX465319 JX421013
Chapsa indica Parnmen018486(RAMK) Thailand JX465295 JX465280
Chapsa leprocarpa GZ19531 China, Guizhou MW009079 MW007981 MW010276
Chapsa leprocarpa GZ19537 China, Guizhou MW009077 MW007984 MW010278
Chapsa leprocarpa GZ19536 China, Guizhou MW009080 MW007982 MW010274
Chapsa niveocarpa HN19508 China, Hainan MW009076 MW010272
Chapsa niveocarpa Lumbsch_19125k2(F) & Mangold (F) Australia, Queensland EU675274
Chapsa niveocarpa Lumbsch 19151p & Mangold (F) Australia, Queensland FJ708487 EU075567
Chapsa patens FJ19131 China, Fujian MT995055 MW007979 MW010275
Chapsa patens FJ19049 China, Fujian MW007918 MW007980
Chapsa wolseleyana FJ19158 China, Fujian MW009078 MW010273 MW010277
Chapsa wolseleyana FJ19148 China, Fujian MW009106 MW010270 MW010279
Chapsa murioelongata HN19222 China, Hainan MW009102 MW010271
Chapsa murioelongata HN19682 China, Hainan MW009103 MW010269
Chapsa pulchra CHAPUL19129t Australia KC020261 KC020255
Astrochapsa meridensis Luecking 17770 (F) Costa Rica EU075655 EU075610
Astrochapsa mastersonii Lumbsch 20500f Fiji JX420996
Astrochapsa zahlbruckneri Papong 6516 Thailand JX421467
Astrochapsa astroidea Lumbsch 19166n & Mangold(F) Australia, Queensland EU075614 EU075566
Astrochapsa astroidea Lumbsch 19750a Thailand JX421441 JX420974
Astrochapsa astroidea Papong 6004 Thailand JX421442 JX420975
Astrochapsa astroidea Luecking 24006 Thailand JX421443 JX420977
Astrochapsa astroidea Luecking 24008 Thailand JX421444 JX420978
Astrochapsa astroidea Luecking 24011 Thailand JX421445 JX465278
Chroodiscus coccineus Herb. R. Luecking 2000 Costa Rica AF465441

Multi-locus (ITS, mtSSU and nuLSU) phylogenetic analysis was performed. The combined analysis included 70 sequences (Table 1) representing 18 in-group taxa and one out-group taxon. As many species as possible of Chapsa s. lat. were contained in our data matrix including the taxa that were similar in morphology or sequence to the new species and the two records. We blasted sequences of the three species in GenBank and selected sequence-similar taxa on a pre-determined cut-off.

The alignment was undertaken by applying MAFFT 7 with the option of L-INS-I (Katoh and Standley 2013). The three single-locus alignments were concatenated in PhyloSuite v1.2.2 (Zhang et al. 2020). The concatenated data matrix comprised 3188 nucleotide sites (nuLSU 1405 bp, ITS 647 bp and mtSSU 1136 bp). In order to check the consistency between the three loci, incongruence length difference test (ILD Test) was carried out using PAUP. The P value of ILD Test was 0.65 (>0.5), so the three loci were suitable for polygenic phylogeny. Construction of the ML (Maximum Likelihood) tree was undertaken by applying RAxML v.8.2.12 (Stamatakis 2014) and implementing a GTRGAMMA model. For BI (Bayesian Inference) analysis, PartitionFinder 2 (Lanfear et al. 2017) was used to determine the best-fit model for each partition. For the nuLSU region, we used GTR+I+G, for ITS, GTR+G, and for mtSSU, HKY+I+G. BI analysis was performed with MrBayes 3.2.7 (Ronquist et al 2012). Markov Chain Monte Carlo (MCMC) chains were run for 200,000 generations, sampling every 100th generation, at which point, the average standard deviation of split frequencies was 0.001738. ML bootstrap values (BS) ≥ 75% and Bayesian posterior probabilities (PP) ≥ 0.95 were considered as significantly supported.

Results and discussion

The BI and ML trees showed similar topologies and thus, only the BI tree was provided (Fig. 1). The three species were all monophyletic with a high support value: C. murioelongata (100%, 1.00), C. wolseleyana (99%, 1.00) and C. niveocarpa (91%, 1.00). Chapsa murioelongata is sister to the clade consisting of C. wolseleyana and C. patens (Nyl.) Frisch. Chapsa niveocarpa HN19508 and C. niveocarpa Lumbsch form a well-supported clade and are sisters to C. leprocarpa.

Figure 1. 

Bayesian phylogenetic tree generated from analysis of combined ITS, nuLSU and mtSSU. Chroodiscus coccineus is the out-group taxon. ML-bootstrap values/Bayesian posterior probabilities above 50% are written next to nodes.


New species

Chapsa murioelongata M.Z. Dou & M. Li, sp. nov.

Figure 2


The specific epithet murioelongata refers to the elongate, muriform ascospores.


China. Hainan Province: Ledong County, Jianfengling National Forest Park, 18°42'39"N, 108°52'37"E, alt. 760 m, on bark, 09 Dec 2019, Y. H. Ju HN19222 (LCUF: holotype: HN19222; GenBank MW009102 for ITS and MW010271 for LSU).


Thallus corticolous, crustose, olive-grey, surface dull, smooth to uneven, ecorticate. Apothecia erumpent, dispersed or two to four aggregated, rounded, 1–3 mm diam.; thalline margin lobed with white felt-like inner surface, lobes strongly backward curved; disc flesh-coloured, covered by thick, white pruina. Exciple 80–105 μm wide laterally, dark brown; epihymenium 20–40 μm high, with coarse greyish granules; hymenium clear, 110–170 μm high, non-amyloid; hypothecium colourless, 10–30 μm high; paraphyses simple, tips unbranched; periphysoides present, 5–30 μm long. Asci 4–6 (8)-spored, clavate, 100–120 × 35–50 μm; ascospores hyaline, bacillary with rounded to subacute ends, submuriform with 20–25 transverse septa and 0–2 longitudinal septa per segment, 75–105 × 9.5–16 μm, non-halonate, I-. Pycnidia not observed.

Figure 2. 

Chapsa murioelongata (LCUF HN19222) A habit of thallus with apothecia at different developmental stages B apothecium (the pruina of the disc partly scraped off) C section of apothecium with periphysoids (direction of arrow) D paraphyses E an ascus containing six ascospores F ascospore. Scale bars: 3 mm (A); 0.5 mm (B); 50 μm (C); 8 μm (D); 30 μm (E); 25 μm (F).


Thallus K-, C-, PD-; no compounds detectable by TLC.

Ecology and distribution

On the bark in semi-exposed forest of Hainan Province.

Additional specimens examined

China. Hainan Province: Changjiang County, Bawangling Nature Reserve, Yajia Scenic Area, 10°04'54"N, 109°07'04"E, alt. 810 m, on bark, 08 Dec 2019, Y. H. Ju HN19167 (LCUF); China. Hainan Province: Lingshui County, Diaoluo Mountain, 18°43'35"N, 109°52'02"E, alt. 900 m, on bark, 14 Dec 2019, M. Li HN19682 (LCUF) (GenBank MW009103 for ITS and MW010269 for LSU).


Chapsa murioelongata is characterised by its olive-grey thallus; white pruinose discs; distinct periphysoids; clear hymenium; 4–8-spored asci; submuriform ascospores with 20–25 transverse septa and 0–2 longitudinal septa per segment. Chapsa microspora Kalb, C. asteliae (Kantvilas & Vězda) Mangold, Astrochapsa elongata Poengs. & Lumbsch and C. patens are morphologically similar to the new species. Chapsa microspora can be distinguished from C. murioelongata by the smaller apothecia (0.6–1.2 mm diam.), transversely septate and smaller ascospores (7–9 × 4 μm) (Lumbsch et al. 2011). Chapsa asteliae differs in amyloid and shorter ascospores (30–80 μm) (Kantvilas and Vězda 2000; Mangold 2008). Astrochapsa elongata differs from C. murioelongata in having shorter ascospores (40–65 μm) and less longitudinal septa per segment (0–1) (Poengsungnoen et al. 2019). Chapsa patens differs from C. murioelongata chiefly in the single-spored asci and broader ascospores (22–35 μm) (Frisch 2006).

Blast searches of nuLSU sequences indicate Chapsa murioelongata has close affinities with C. patens (98.36% identity), C. wolseleyana (95.63% identity), C. leprocarpa (91.97% identity) and C. indica (90.81% identity), so all these species were included in the phylogenetic analyses. Chapsa murioelongata was well separated from any other species in the tree and strongly supported as the monophyletic (PP = 1; ML = 100%).

New records

Chapsa wolseleyana Weerakoon, Lumbsch & Lücking, in Weerakoon, Rivas Plata, Lumbsch & Lücking, Lichenologist 44(3): 377 (2012)

Figure 3

Astrochapsa wolseleyana (Weerakoon, Lumbsch & Lücking) Parnmen, Lücking & Lumbsch, in Parnmen et al., PLoS ONE 7(12): 10 (2012)


Thallus crustose, corticolous, grey-brown, surface dull to slightly shiny, uneven, fissured. Apothecia erumpent, dispersed, sometimes two or three fused, mostly rounded to seldom slightly angular, 0.7–1.2 mm diam.; thalline margin raised to lobulate, lobes erected to recurved, inner part brown, covered with rose-red or white pruina; disc exposed, rose-red, covered with thick, rose-red pruina. Exciple fused, cupular, laterally 180–250 μm wide, yellowish-brown to brown; epihymenium rose-red with granules, 20–50 μm high, K+ green; hymenium 140–230 μm high, clear, colourless, non-amyloid; hypothecium indistinct; paraphyses septate, tips rose-red and moniliform with oval or rectangular cells; periphysoides present, 50–100 μm long. Asci clavate, 1-spored, 110–135 × 35–50 μm; ascospores densely muriform, oblong-ellipsoid, with hemispherical to roundish ends, 105–130 × 30–45 μm, first reddish, becoming hyaline to slightly olive-brown at maturity, I-. Pycnidia not observed.

Figure 3. 

Chapsa wolseleyana (LCUF FJ19148-b) A habit of thallus with apothecia B apothecia at different developmental stages C apothecium (part of pruina scraped off) D section of apothecium with periphysoids (direction of arrow) E paraphyses F young and mature ascospores. Scale bars: 1.5 mm (A); 1 mm (B); 0.25 mm (C); 120 μm (D); 10 μm (E); 25 μm (F).


No substances detected by TLC but apothecial disc with pigment producing K+ yellow-green efflux, suggesting presence of isohypocrelline.

Ecology and distribution

Growing on bark exposed to wind and high light intensity in montane forests. Worldwide distribution: Sri Lanka (Weerakoon et al. 2012) and newly reported for China.

Selected specimens examined

China. Fujian Province: Quanzhou City, Jiuxian Mountain, Reflecting Pool, 25°42'57"N, 118°07'14"E, alt. 1540 m, on bark, 5 Jul 2019, F.Y. Liu FJ19148-b (LCUF) (GenBank MW009106 for ITS, MW010270 for LSU and MW010279 for SSU); China. Fujian Province: Quanzhou City, Jiuxian Mountain, Natural Observation Path, 25°42'44"N, 118°07'17"E, alt. 1460 m, on bark, 25 Jul 2019, F.Y. Liu FJ19158 (LCUF) (GenBank MW009078 for ITS, MW010273 for LSU and MW010277 for SSU). China. Fujian Province: Quanzhou City, Jiuxian Mountain, Reflecting Pool, 25°42'57"N, 118°07'14"E, alt. 1540 m, on bark, 25 Jul 2019, F.Y. Liu FJ19127-2, same locality, FJ19128-2, FJ19141-2 (LCUF).


Chapsa wolseleyana is characterised by its grey-brown, uneven thallus, apothecia with raised to lobed thalline margin, rose-red discs with similar coloured pruina, rose-red epihymenium and paraphyses tips, distinct periphysoids, 1-spored asci, muriform ascospores, red when young and hyaline to olive-brown when old. Only a few species of Chapsa have pigmented discs and among them C. rubropulveracea Hale ex Mangold, Lücking & Lumbsch is morphologically most similar to C. wolseleyana, but its thallus is farinose and its ascospores are 8 per ascus, smaller (15–20 ×5–6 μm) and transversely septate (Lumbsch et al. 2011).

Chapsa wolseleyana was transferred to Astrochapsa, based on a phenotype-based analysis (not molecular phylogeny) (Parnmen et al. 2012). However, our phylogenetic analysis shows that this species belongs in Chapsa, rather than Astrochapsa. Chapsa wolseleyana was associated phylogenetically with a strongly-supported clade (100/1) with C. patens, but with sufficient distance to be considered a distinct species. In addition, the latter differs from C. wolseleyana in having larger pale brown apothecia (up to 2 mm diam.) with white pruina, unpigmented epihymenium and unpigmented paraphyses adspersed with fine greyish to brownish granules, hyaline ascospores (Frisch et al. 2006; Joshi et al. 2012; Joshi et al. 2018).

Chapsa niveocarpa Mangold in Mangold, Elix & Lumbsch, Flora of Australia, 57:654 (2009)

Figure 4


Thallus corticolous, crustose, pale grayish-green surface dull and fluctuating along the bark. Apothecia erumpent, solitary to fused, angular rounded to slightly elongate, 0.5–1.8 × 0.5–1.2 mm; thalline margin split and recurved, insidewith thick white pruina; disc exposed, yellowish-brown, covered by white pruina. Exciple laterally 12–75 μm wide, dark brown; epihymenium 10–20 μm high; hymenium 120–200 μm high, grey-brown, inspersed by granules, non-amyloid; hypothecium indistinct; paraphyses unbranched; tips distinctly thickened; periphysoides present, but obscured by granular inclusions. Asci 1-spored, clavate, 120–140 × 27–36 μm; ascospores densely muriform, with thick halo at both ends, oblong, hyaline, 115–135 × 25–34 μm, I-. Pycnidia not observed.

Figure 4. 

Chapsa niveocarpa (LCUF HN19508) A habit of thallus with apothecia B apothecium (part of pruina scraped off) C section of apothecium with periphysoids (direction of arrow) D paraphyses with hyaline granules E ascus F ascospore with halo. Scale bars: 1 mm (A); 0.5 mm (B); 50 μm (C); 25 μm (D); 30 μm (E); 25 μm (F).


Thallus K-, C-, PD-; no compounds detectable by TLC.

Ecology and distribution

Growing on tree bark in tropical rainforests in altitudes ranging from 500 to 1100 m. Australia, Queensland (Mangold 2008); newly reported for China.

Selected specimens examined

China. Hainan Province: Wuzhishan City, Wuzhishan Nature Reserve, 18°54'13"N, 109°41'04"E, alt. 870 m, on bark, 12 Dec 2019, M. Li HN19508 (LCUF) (GenBank MW009076 for ITS and MW010272 for LSU); China. Hainan Province: Wuzhishan City, Wuzhishan Nature Reserve, 18°53'13"N, 109°41'04"E, alt. 1020 m, on bark, 12 Dec 2019, M. Li HN19530 (LCUF); China. Hainan Province: Wuzhishan City, Wuzhishan Nature Reserve, 18°54'13'N, 109°41'04'E, alt. 870 m, on bark, 12 Dec 2019, M. Li HN19499 (LCUF); China. Hainan Province: Lingshui County, Diaoluo Mountain, 18°43'35"N, 109°52'02"E, alt. 900 m, on bark, 14 Dec 2019, M. Li HN19687 (LCUF); China. Hainan Province: Lingshui County, Diaoluo Mountain, 18°43'35"N, 109°52'02"E, alt. 900 m, on bark, 14 Dec 2019, M. Li HN19679 (LCUF).


Chapsa niveocarpa is characterised by its crustose, pale greyish-green thallus; rounded to elongate apothecia, yellowish-brown discs with white pruina, obscured periphysoids, inspersed hymenium, 1-spored(rare 2-spored)ascus and muriform and hyalineascospores with halo. Chapsa niveocarpa is morphologically similar and phylogenetically related to C. leprocarpa, and both species occur on bark in tropical forests (Frisch 2006; Mangold 2008; Parnmen et al. 2012). Chapsa leprocarpa differs from C. niveocarpa in having a lower hymenium (100–130 μm) and smaller ascospores (up to 111 μm long) (Frisch 2006). The specimen (HN19508) we collected in China is allocated phylogenetically to a strongly-supported (1/91) clade with C. niveocarpa. The collections cited above are the first reports for China.

Key to Chapsa in China

1 Disc with red pruina; ascospores 1/ascus, muriform, 105–135 × 30–50 μm C. wolseleyana
Disc with white pruina 2
2 Ascospores transversely septate; ascospores 4–8/ascus, 50–110 × 6–12 µm C. indica
Ascospores (sub)muriform 3
3 Hamathecium inspersed; ascospores 1/ascus, 80–190 × 20–50 μm 4
Hamathecium clear 5
4 Ascospores 1/ascus, 80–190 × 20–50 μm C. niveocarpa
Ascospores 8/ascus, 40–50 × 11–15 μm C. mirabilis
5 Asci 4‒6 (8)-spored; acsospores oblong to cylindrical with rounded to subacute ends, submuriform with 20–25 transverse septa and 0–2 longitudinal septa per segment, 75–105 × 9.5‒16 μm C. murioelongata
Asci 4-spored; acsospores oblong to slightly ellipsoid, with roundish ends, 60–130 × 20–40 μm C. leprocarpa


This study was supported by the National Natural Science Foundation of China (31750001), Doctoral Initiation Fund of Liaocheng University (318051813) and Research Fund of Liaocheng University (318012011).


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1 *These two authors contributed equally to this work.
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