Genome sequencing of Phycotheca Boreali-Americana material of Ceramothamnion codii H.Richards confirms synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards

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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / http://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (électronique / electronic) : 1776-0984 Directeur De la publication / Publication director: Gilles BLOCH Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief: Line le Gall Muséum national d’Histoire naturelle assistant De réDaction / assistant editor: Violette GRUNENBERGER ([email protected]) Mise en paGe / Page layout: Violette GRUNENBERGER réDacteurs associés / associate editors Ecoevolutionary dynamics of algae in a changing world Stacy KRUEGER-HADFIELD Virginia Institute of Marine Science Eastern Shore Laboratory, Wachapreague, VA 23480 (United States) Jana KULICHOVA Department of Botany, Charles University, Prague (Czech Republic) Cecilia TOTTI Dipartimento di Scienze della Vita e dell’Ambiente, Università Politecnica delle Marche, Via Brecce Bianche, 60131 Ancona (Italy) Phylogenetic systematics, species delimitation & genetics of speciation Sylvain FAUGERON UMI3614 Evolutionary Biology and Ecology of Algae, Departamento de Ecología, Facultad de Ciencias Biologicas, Pontificia Universidad Catolica de Chile, Av. Bernardo O’Higgins 340, Santiago (Chile) Marie-Laure GUILLEMIN Instituto de Ciencias Ambientales y Evolutivas, Universidad Austral de Chile, Valdivia (Chile) Diana SARNO Department of Integrative Marine Ecology, Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Napoli (Italy) Comparative evolutionary genomics of algae Nicolas BLOUIN Department of Molecular Biology, University of Wyoming, Dept. 3944, 1000 E University Ave, Laramie, WY 82071 (United States) Heroen VERBRUGGEN School of BioSciences, University of Melbourne, Victoria, 3010 (Australia) Algal physiology & photosynthesis Janet KÜBLER California State University Northridge, Department of Biology, California State University, Northridge, CA 91330-8303 (United States) Prokaryotic algae Nico SALMASO IASMA Research and Innovation Centre, Fondazione Mach-Istituto Agrario di S. Michele all’Adige, Limnology and River Ecology, Via E. Mach, 1, 38010 San Michele all’Adige, Trento (Italy) Vitor VASCONCELOS Faculdade de Ciências da Universidade do Porto and CIIMAR, Rua do Campo Alegre, s/n, 4169-007 Porto (Portugal) couverture / cover: Extrait de la Figure 2/Extract of Figure 2 89 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.cryptogamie.com/algologie Genome sequencing of Phycotheca Boreali-Americana material of Ceramothamnion codii H.Richards confirms synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards Maria Beatriz BARROS-BARRETO Department of Botany, Federal University of Rio de Janeiro, Rua Professor Rodolpho P. Rocco 211, CCS-Bloco A, sala 66, 21941-902, Rio de Janeiro, RJ (Brazil) [email protected] (corresponding author) Jeffery R. HUGHEY Division of Mathematics, Science, and Engineering Hartnell College, CA 93901, Salinas, California (United States) Christine A. MAGGS School of Biological Sciences, Queen’s University, Belfast BT9 5AJ (North Ireland) Submitted on 5 August 2024 | Accepted on 1 December 2024 | Published on 15 December 2025 Barros-Barreto M. B., Hughey J. R. & Maggs C. A. 2025. — Genome sequencing of Phycotheca Boreali-Americana material of Ceramothamnion codii H.Richards confirms synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards. Cryptogamie, Algologie 46 (7): 89-113. https://doi.org/10.5252/cryptogamie-algologie2025v46a7. http://cryptogamie.com/algologie/46/7 ABSTRACT The systematic status of the newly proposed genus Stirkia Barros-Barreto & Maggs and its relationship to the currently accepted name Ceramothamnion H.Richards remain uncertain because no authentic specimens of the generitype C. codii H.Richards from the type locality in Bermuda have been sequenced. To clarify the taxonomy of C. codii, we performed high-throughput sequencing on a topotype, a 1912 Phycotheca Boreali-Americana (P.B.-A.) specimen collected by F.S. Collins. The rbcL, COI-5P and LSU genetic markers from the P.B.-A. specimen were assembled and used to infer gene and multi-gene phylogenies to determine its phylogenetic history. In all analyses, the topotype of C. codii was closely allied with C. fujianum Barros-Barreto & Maggs, the generitype of Stirkia, as well as with other species of Ceramothamnion from the southwestern Atlantic and Pacific Oceans. These data support the relegation of the genus Stirkia to Ceramothamnion and indicate that the only previously published DNA sequences of C. codii, from Hawai‘i, represent a different, presumably undescribed species of Ceramothamnion. They also highlight the power of whole genome sequencing and the need to analyse type, topotype, or historically authentic herbarium specimens to ensure the accurate application of algal binomials. RÉSUMÉ Le séquençage du génome du matériel Phycotheca Boreali-Americana de Ceramothamnion codii H.Richards confirme la synonymie de Stirkia Barros-Barreto & Maggs avec Ceramothamnion H.Richards. Le statut systématique du genre Stirkia Barros-Barreto & Maggs nouvellement proposé et sa relation avec le nom actuellement accepté Ceramothamnion H.Richards restent incertains car aucun spécimen authentique du généritype C. codii H.Richards de la localité type des Bermudes n’a été séquencé. Pour clarifier la taxonomie de C. codii, nous avons effectué un séquençage à haut débit sur le spécimen d’un topotype, un spécimen de Phycotheca Boreali-Americana (P.B.-A.) de 1912 collecté par F.S. Collins. Les marqueurs génétiques rbcL, COI-5P et LSU du P.B.-A. des spécimens ont été assemblés et utilisés pour déduire des phylogénies génétiques et multigéniques afin de déterminer leur histoire KEY WORDS Ceramieae, COI-5P, LSU, molecular systematics, rbcL gene. 90 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. phylogénétique. Dans toutes les analyses, le topotype de C. codii était étroitement lié à C. fujianum Barros-Barreto & Maggs, le généritype de Stirkia, ainsi qu’à d’autres espèces de Ceramothamnion du sud-ouest des océans Atlantique et Pacifique. Ces données soutiennent la relégation du genre Stirkia à Ceramothamnion et indiquent que les seules séquences d’ADN précédemment publiées de C. codii, d’Hawaï, représentent une espèce différente et non décrite de Ceramothamnion. Ils mettent également en évidence la puissance du séquençage du génome entier et la nécessité d’analyser des spécimens d’herbier de type, de topotype ou historiquement authentiques pour garantir l’application précise des binômes végétaux. MOTS CLÉS Ceramieae, COI-5P, LSU, systématique moléculaire, rbcL gène. INTRODUCTION Recently, an extensive morphological and molecular reclassification of the tribe Ceramieae Roth was proposed (Barros-Barreto et al. 2023a). The new genus Stirkia Barros-Barreto & Maggs was described for a morphologically diverse clade identified by its well-supported molecular monophyly, typified by Stirkia fujiana Barros-Barreto & Maggs. Based on a partial LSU rDNA sequence of “Ceramium codii” from Kure Atoll, Hawai‘i (HQ421967; “C. codii” was included in Stirkia and the new combination S. codii (H.Richards) Barros-Barreto & Maggs was proposed (Barros-Barreto et al. 2023b). Wynne & Schneider (2023) promptly noted that the basionym of Ceramium codii was Ceramothamnion codii H.Richards, a species originally described from specimens forming minute red turfs that grew epiphytically on Codium tomentosum Stackhouse from Bermuda (Richards 1901). As Ceramothamnion H.Richards is a valid genus with nomenclatural priority, Wynne & Schneider (2023) transferred all the species assigned to Stirkia by Barros-Barreto & Maggs to Ceramothamnion. They noted however that if additional phylogenetic evidence demonstrates that C. codii is phylogenetically distinct from S. fujiiana, Stirkia would remain available for use. To determine the correct application of a name, analysing DNA from the type material is preferable (Hughey et al. 2001). As documented by Wynne & Schneider (2023), the type material of Ceramothamnion codii Richards (1901: 265) is no. 845 in Phycotheca Boreali-Americana (P.B.-A.) Fasc. XVII (Collins et al. 1901, coll. H.M. Richards, Harris Bay, Bermuda, Feb. 1889, on Codium tomentosum [actually C. decorticatum (Woodward) M.Howe, see Schneider 2003]). However, it is not possible to analyse the BM (herbarium acronyms follow Thiers 2024, continuously updated) isotype of C. codii as the specimen distributed as no. 845 (BM000619304) is mounted on mica and has developed a fungal growth that has destroyed the algal material and rendered it demonstrably ambiguous (Fig. 1A). We do not have access to the NY specimens and thus performed high-throughput sequencing on another P.B.-A. specimen of C. codii in BM (Fig. 1B, BM000936098), a specimen collected by F.S. Collins from Cooper’s Island, Bermuda, distributed in the exsiccata P.B.-A. as no. 1899, by Collins et al. (1912) and considered topotype material by Cho & Fredericq (2006) and Huisman (2018). The lectotype of C. codii has never been formally designated, despite the assumption that it should be the NY example (Huisman 2018). The assembled genetic markers that we obtained were then used to perform single and multi-marker phylogenetic analyses to determine the systematic relationship of C. codii to S. fujiiana and the other species recently transferred to Ceramothamnion. MATERIAL AND METHODS DNA extrActioN AND geNomic ANAlysis Epiphytic fragments of the tiny filamentous red alga distributed as Ceramothamnion codii P.B.-A. no. 1899 were sampled from the left and right specimens of Codium sp. (BM000936098; Fig. 1) and combined for the DNA extraction. The sample was processed in the personal residence of JRH. Genomic DNA was isolated using the DNeasy Blood and Tissue Kit (Qiagen) following previous protocols and two modifications: the binding step was centrifuged at 4 000 g for 3 mins and the DNA was eluted after incubation for 7 mins in 40 µL TAE (Hughey & Gabrielson 2012, Lindstrom et al. 2011, Hughey et al. 2019). The 150 bp PE library was constructed with the KAPA HyperPrep Kit (Roche, Basel, Switzerland) and sequenced by Quick Biology Inc. (Pasadena, California, United States). The resulting reads were filtered using the default BBDuk settings in Geneious Prime 2019.1.3 (Biomatters Limited). The filtering yielded 49 408 547 trimmed reads that were assembled de novo with MEGAHIT 1.2.9 using the default settings with the kmers 29, 39, 59, 79, 99, 119, 141 (Li et al. 2015). Since the de novo did not yield long algal contigs, the standard red algal genetic markers (COI, rbcL, LSU) were instead assembled by iteratively mapping reads 5X onto the following Campylaephora sungminbooi (J.R.Hughey & G.H.Boo) Barros-Barreto & Maggs reference sequences using the Medium Sensitivity/ Fast setting in Geneious Prime 2019.2.3: rbcL-GenBank accession number KR025491; COIGenBank accession number KU145004; and LSU-GenBank accession number EF033604. The assembly sequences were then confirmed for accuracy by iterative remapping of the reads onto the consensus sequences using the same settings in Geneious Prime. Codium rbcL and tufA DNA sequences were also targeted using the same assembly parameters above using the Codium reference sequences: rbcL-GenBank accession number ON113499, tufA-GenBank accession number 91 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) KX855797, and rpl16-rps3-GenBank accession number EU045560 to confirm the identity of the host. PhylogeNetic ANAlyses The Ceramothamnion codii marker sequences determined here and others downloaded from GenBank (Appendix 1) were aligned using Seaview 4.2.10 (Gouy et al. 2010). There were totals of 182 accessions (including the outgroup) for the rbcL locus; 84 accessions for COI-5P; 87 for LSU rDNA; and 72 for the concatenated alignment of the three markers (Appendix 2). Due to missing marker data for some taxa, the rbcL alignment was trimmed to 1239 bp and the COI-5P alignment to 624 bp. For LSU, ambiguous sites were cut according to GBlocks (Castresana 2000) to 573 bp following the alignment with MAFFT v7 (Katoh et al. 2019); the concatenated alignment was 2487 bp (rbcL = 1239 bp; LSU = 682 bp; COI-5P = 566 bp). Sequence data were analysed with the maximum likelihood (ML) optimality criterion using IQ-TREE (Trifinopoulos et al. 2016; Nguyen et al. 2015). Genes and codons were partitioned, and evolutionary models were selected by ModelFinder (Kalyaanamoorthy et al. 2017) using the Bayesian Information Criterion (BIC); partitioned models were used when appropriate (Chernomor et al. 2016); merge option was also included to increase model fit in IQ-TREE (Appendix 2). Tree branches were tested by SH-like aLRT (Guindon et al. 2010), Ultrafast (Hoang et al. 2018) and standard bootstrapping with 1000 replicates. Bayesian inference (BI) analyses were implemented using MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001; Ronquist & Huelsenbeck 2003) or executed on the CIPRES Science Gateway (Miller et al. 2010). BI analyses consisted of two parallel runs, each of four Markov chains, and 306 generations were run (three times for each dataset) with sampling every 1000 generations. The rbcL and COI-5P markers were partitioned (codon positions), the concatenated alignment was partitioned by gene and codon, and the model implemented independently for each gene/codon partition; all the parameters were unlinked. Burn-in fraction was set to 0.30%, and the generation numbers stopped when the Average Standard Deviation of split frequencies reached 0.015 (Appendix 2). The full taxon rbcL and COI-5P trees were fiG. 1. — Ceramothamnion codii H.Richards type and authentic material: A, C. codii, from Cooper’s Island, Bermuda in the exsiccata Phycotheca Boreali-Americana (P.B.-A.) no. 845 (BM000619304) on Codium tomentosum Stackhouse (actually C. decorticatum (Woodward) M.Howe)); B, C. codii, from Cooper’s Island, Bermuda in the exsiccata Phycotheca Boreali-Americana (P.B.-A.) no. 1899 (BM000936098). Scale bars: A1, B1, 1 cm; A2, B2, 1 mm. AA1A2 BB1 B2 92 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. rooted designating five species from Callithamniaceae and Wrangeliaceae, as the outgroups. For the LSU analyses, two species of Ceramiaceae were used as outgroups. Antithamnion nipponicum Yamada & Inagaki (Ceramiaceae, Antithamnieae) was selected as the outgroup for the concatenated alignment (Appendix 2). fiG. 2. — Ceramothamnion codii H.Richards growing on Codium tomentosum Stackhouse and Codium decorticatum (Woodward) M.A.Howe thalli, and original drawings from the Algae of Bermuda (Collins & Hervey 1917), lectotype NY02116306. Scale bar: 10 cm. 93 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) RESULTS We herein designate Ceramothamnion codii no. 845 P.B.-A. (NY02116306) collected by H. M. Richards from Harris Bay, Bermuda, February, 1898 as the lectotype (Fig. 2), on the basis that Collins’ herbarium is in NY (Setchell 1925). Ceramothamnion codii no. 1899 P.B.-A. (BM000936098) collected by F.S. Collins from Cooper’s Bay, Bermuda, April 29, 1912 is considered to be a topotype (Fig. 1). High-throughput sequencing of the BM’s P.B.-A. no. 1899 Ceramothamnion codii specimen yielded 53 767 172 reads (BioSample SAMN41642873, BioProject ID PRJNA1119217, SRA SRR29258986). De novo assembly of the trimmed reads resulted in 1 338 contigs with an N50 of 418 bp and maximum C. codii contig length of 1 920 bp, which was inadequate for organellar genome assembly. Mapping of the reads to the reference sequences however resulted in the assembly of the complete C. codii rbcL (GenBank accession number PP868167, 1467 bp) and COI (GenBank accession number PP868169, 1584 bp) genes, and a partial nuclear encoded LSU sequence (GenBank accession number PP863326, 1585 bp) with 144X, 31X, and 13X coverages. Phylogenetic analysis of the complete rbcL gene sequence of Ceramothamnion codii positioned it in a clade containing species recently reclassified as Ceramothamnion from the southwestern Atlantic, and western (including C. fujianum) and eastern Pacific Oceans (Fig. 3). The clade containing C. codii and various Ceramothamnion spp. was well-supported based on Bayesian and ultrafast bootstrap numbers, however, it displayed low support based on standard bootstrapping (Fig. 3), the genetic distance among this clade is 7-10% (Appendix 3 , Table 1). The LSU analysis placed C. codii close to C. fujianum, however the inferred tree indicated a paraphyletic relationship with respect to C. codii and the many species currently assigned to Ceramothamnion (Fig. 4), the genetic divergence from C. codii from Ceramothamnion spp. is 3-7%, except from C. brasiliense (16%) (Appendix 3, Table 2). Analysis of the COI-5P data resulted in a phylogenetic tree with numerous polytomies. Ceramothamnion codii was grouped in an unsupported clade with six Ceramothamnion spp. (Fig. 5; Appendix 3, Table 3). The combined analysis fully resolved C. codii in a clade with C. fujianum and eleven other Ceramothamnion spp. (Fig. 6). Further analysis of the genome sequencing data found that the LSU sequence identified in GenBank as “Ceramium codii” from Kure Atoll, Hawai‘i differed from the Ceramothamnion codii topotype from Bermuda by 3.7%. In addition, mapping of the genome data to rbcL and tufA sequences of Codium tomentosum failed to yield matching sequences. Instead, the two Codium specimens on sheet fiG. 3. — Phylogenetic reconstruction of the red algal tribe Ceramieae C.Agardh ex Greville inferred from rbcL gene sequences. The tree is the majority rule consensus tree resulting from Bayesian analysis. Values at the nodes represent posterior probability and bootstrap values for ML, support lower than 0.90 (PP), 80 (Ultrafast Bootstrap) and 40 (standard BP) are not shown. *, full support. Sequence generated in this study is in bold and the topotype sequence is underlined. Reinboldiella Carpoblepharis Herpochondria Ceramium Campylaephora Ceramieae sp.1 MW366525 Pseudoceramium Yoneshiguea Ceramieae sp.2 MK185813 Ceramothamnion pacificum FJ795539 Ceramothamnion pacificum GQ179817 Ceramothamnion sinicola AY155531 Ceramothamnion sinicola GQ179818 Ceramothamnion codicola AY155522 Ceramothamnion interruptum AY155527 Ceramothamnion riosmenae JN712693 Ceramothamnion horridulum AF521796 Ceramothamnion coulteri MW366528 Ceramothamnion coulteri GQ252482 Ceramothamnion sp. 1 MW366508 Ceramothamnion vagans MW366529 Ceramothamnion sp. 2 MW366530 Ceramothamnion dumosertum GQ252454 Ceramothamnion japonicum A Y948753 Ceramothamnion fujianum DO458949 Ceramothamnion fujianum DO458952 Ceramothamnion inconspicuum LR031255 Ceramothamnion julieae MW366506 Ceramothamnion codii PBA1899 Bermuda PP868167 Ceramothamnion brasiliense DO458922 Ceramothamnion sp. 3 KF367768 Celeceras Gayliella Ceramieae sp. 4 MW366513 Ceramieae sp. 3 GQ252468 Corallophila atlantica MW366512 Corallophila atlantica MW366536 Ceramieae sp. 5 DO458953 Corallophila eatoniana Ceramieae sp. 6 MW366511 Centroceras Spermothamnion repens U04024 Sphondylothamnion multifidum AF439312 Spyridia filamentosa AF458705 Aglaothamnion tenuissimum MW366516 Seirospora interrupta AF43931 0 1.00/100/75 1.00/100/81 1.00/100/87 1.00/100/70 1.00/100/- 1.00/81/- -/85/- 1/100/99 1.00/-/- 1.00/100/99 0.99/100/78 0.99/100/87 0.99/100/93 1.00/100/87 1.00/100/94 -/100/85 0.94/100/- 1.00/100/- 1.00/100/81 1.00/100/43 0.97/-/- 1.00/98/72 0.91/98/72 1.00/100/99 1.00/100/95 0.99/98/- 1.00/97/- 1.00/89/- 1.00/100/99 1/-/- 1.00/89/- 0.91/83/- 1.00/99/86 1.00/99/98 Ceramothamnion 0.03 * * * * * * * * 94 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. no. 1899 P.B.-A. yielded rbcL and tufA sequences that were identical to sequences deposited in GenBank under the names Codium decorticatum (Woodward) M.A.Howe and Codium fragile. The rpl16-rps3 sequence of Codium fragile was a partial sequence and did not span the entire locus, but did contain nucleotide markers G at site 50, C at 104, and C at 134. These markers are identical to Codium fragile subsp. tomentosoides and the Codium fragile haplotype from China. DISCUSSION cerAmothAmNioN coDii genetic analysis The rbcL gene is the most commonly used marker within the Ceramiales (Cassano et al. 2019; Díaz-Tapia & Verbruggen 2024), therefore obtaining an rbcL sequence from authentic material of Ceramothamnion codii made it possible to phylogenetically confirm the circumscription of Ceramothamnion. High-throughput sequencing of red algal type and other fiG. 4. — Phylogenetic reconstruction of the red algal tribe Ceramieae C.Agardh ex Greville inferred from LSU rDNA sequences. The tree is the majority rule consensus tree resulting from Bayesian analysis. Values at the nodes represent posterior probability and bootstrap values for ML, support lower than 0.90 (PP), 80 (Ultrafast Bootstrap) and 40 (standard BP) are not shown. *, full support. Sequence generated in this study is in bold and the topotype sequence is underlined. Abbreviations: AU, Australia; BE, Bermuda; BC MX, Baja California, Mexico; CA US, California; United States; CH, China; CL, Chile; FL US, Florida, United States; HI US, Hawaiʻi, United States; IR, Ireland; IT, Italy; JA, Japan; KO, South Korea; MX, Mexico; NS CA, Nova Scotia, Canada; OR US, Oregon, United States; PA, Panama; PE, Peru; SA, South Africa; SP, Spain; TX US, Texas, United States. Centroceras gasparrinii BR MW354751 Centroceras gasparrinii BR D0374349 Centroceras gasparrinii BR MN 105062 Centroceras gasparrinii BR MN 105062 Centroceras gasparrinii HI US HQ422061 Centroceras gasparrinii HI US HQ422191 Centroceras gasparrinii HI HQ422101 Centroceras gasparrinii HI US HQ422498 Centroceras gasparrinii BR MW354752 Centroceras gasparrinii HI US D0374354 Centroceras gasparrinii PA D0374359 Centroceras gasparrinii FL US D037 4346 Centroceras gasparrinii TX US D037 4345 Centroceras hyalacanthum FLUS D037 4366 Centroceras hyalacanthum FLUS D037 4367 Centroceras hyalacanthum HIUS HQ422526 Centroceras tetrachotomum SA D037 4360 Centroceras clavulatum PE D0374363 Centroceras clavulatum D0238801 Centroceras clavulatum CA US D037 4362 Centroceras clavulatum CA US D0374361 Centroceras micracanthum PA D037 4365 Gayliella sp. 6 HI US HQ422107 Gayliella jolyana BR MN105060 Gayliella taylorii CA US D0238812 Gayliella jolyana BR MN105061 Ceramieae sp. 4 BR MW354757 Gayliella sp. 9 HI US HQ422195 Gayliella tamoiensis BR MN105056 Gayliella tamoiensis BR MN105055 Gayliella womersleyi AU D0238808 Gayliella sp. 8 HI US HQ422171 Gayliella ardissonei BR MN105050 Gayliella transversalis FL US D0238810 Gayliella mazoyerae IT D0238809 Gayliella sp. 5 BR MN105051 Gayliella iemanja BR MN105052 Gayliella sp. 7 HI US HQ422193 Gayliella sp. 4 BR MN105054 Gayliella dawsonii BR MN105057 Corallophila sp. HI US HQ422146 Corallophila eatoniana WA US D02388022 Ceramieae sp. 5 BR MW354754 Pseudoceramium tenerrimum (Cer. tenerrimum) KO D0238805 Ceramieae sp. 17 HI US HQ421931 Ceramieae sp. 15 HI US HQ422004 Ceramieae sp. 16 HI US HQ422033 Ceramieae sp. 18 HI US HQ422196 Ceramieae sp. 20 HI US HQ422173 Ceramieae sp. 8 (Cer. hanaense) HI US HQ421934 Pseudoceramium oliveirarum (Cer. brevizonatum var. caraibicum) BR MW354756 Campylaephora kondoi (Cer. kondoi) KO D0238804 Campylaephora kondoi (Cer. kondoi) CH KC795847 Ceramieae sp. 19 HI US HQ421961 Campylaephora borealis KO D0238798 Ceramieae sp. 13 (Ceramium sp_.) HI US HQ422054 Campylaephora hypnaeoides JA D0238797 Pseudoceramium tenerrimum (Cer. tenerrimum) CH KC795867 Ceramieae sp. 10 (Cer. diaphanum) CH KC795860 Ceramieae sp. 14 (Ceramium sp.) HI US HQ421687 Ceramieae sp. 7 (Cer. womersfeyi) HI US HQ422536 Yoneshiguea affinis (Cer. affine) MX D0238803 Ceramieae sp. 21 HI US HQ422110 Ceramothamnion sp. 4 HI US HQ422441 Ceramothamnion dumosertum HI US HQ421685 Ceramothamnion sp. 5 HI US HQ422096 Ceramothamnion sp. HI US HQ421967 Ceramothamnion brasiliensis BR MW354747 Ceramiothamnion japonicum CH KC795870 Ceramothamnion codicola OR US D0238806 Ceramothamnion horridulum BC MX D0238807 Centroceras rodmanii CL D037 4364 Ceramothamnion fujianum BR MW354759 Ceramothamnion codii PBA 1899 BE PP863326 Herpochondria borealis (M. borealis) KO D0238799 Herpochondria elegans KO D0238800 Ceramium botryocarpum IR MW354748 Ceramium botryocarpum IR PQ047457 Ceramium virgatum NS CA EF033604 Ceramium pallidum IR PQ047546 Ceramium secundatum IR MW354749 Ceramium gaditanum SP MW354753 Antithamnion nipponicum D0238813 * * * 1.00/98/93 0.94/86/58 1.00/96/76 0.94/96/85 -/99/57 -/100/100 0.96/96/67 -/95/48 -/96/49 -/80/41 1.00/97/60 1.00/97/76 0.90/96/80 1.00/96/83 0.98/94/71 1.00/-/- -/95/- 0.96/98/- 0.94/85/- 0.99/98/- 0.93/-/- -/81/- 0.96/82/- -/89/- 1.00/100/671.00/94/71 -/98/68 0.93/91/64 1.00/99/66 1.00/100/97 1.00/100/99 0.9/87/61 0.96/98/82 -/97/- -/92/58 -/100/100 * * ** * 0.05 95 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) historically valuable herbarium specimens has been shown to be useful for resolving many systematic questions in the red algae (Hughey et al. 2014, Boo et al. 2016, Gabrielson et al. 2018, Hughey & Miller 2021, Hughey et al. 2022). Although whole genome sequencing here was unable to decipher the complete organellar genomes as reported by the studies above, this analysis successfully generated a sufficient number of reads to assemble the standard phylogenetic genetic markers required to conclusively characterize the generic affinity of the 112 year-old topotype C. codii specimen from Bermuda. Although low standard bootstrap supports were observed for the rbcL, LSU and combined datasets, all three analyses positioned Ceramothamnion species together in a single clade. In addition, these analyses show that C. codii is placed in the same clade as the generitype of Stirkia, S. fujiiana, and therefore support the transfer of all Stirkia spp. to Ceramothamnion as proposed by Wynne & Schneider (2023). Seo et al. (2023) resolved a fully supported Ceramothamnion clade in their maximum likelihood phylogeny of the Ceramiales based on nucleotide sequences of 23 protein-coding genes. In addition, our results indicate that the two publicly available sequences in GenBank from Hawai‘i deposited under the name Ceramium codii (collection number ARS03339, LSU and 23S plastid gene sequences) do not represent the same species as the type material from Bermuda. The genetic distances exhibited between the Hawai‘i and Bermuda specimens are higher than fiG. 5. — Bayesian tree inferred from COI-5P sequences. The trees are the majority rule consensus tree resulting from Bayesian analysis. Values at the nodes represent posterior probability and bootstrap values for ML, support lower than 0.90 (PP), 80 (Ultrafast Bootstrap) and 50 (BP) are not shown. *, full support. Sequences generated in this study are in bold and the topotype sequence is underlined. Abbreviations: AU, Australia; BE, Bermuda; BR, Brazil; CA US, California, United States; CH, China; FR, France; GB TR, British Overseas Territory of Tristan da Cunha; IR, Ireland; KO, South Korea; HI US, Hawaiʻi, United States; MA US, Massachusetts, United States; ME US, Maine, United States; NB CA, New Brunswick Canada; NC US, North Carolina; United States; NO, Norway; RI US, Rhode Island, United States; SA, South Africa. Ceramium secundatum IR MW354763 Ceramium secundatum RI US KT250208 Ceramium secundatum FR EU194972 Ceramium secundatum NO MN 184499 Ceramium secundatum NO MN184367 Ceramium secundatum GB TR MK202399 Ceramium secundatum GB TR MK202407 Ceramium sp. 1 NO MN184330 Ceramium virgatum MA US KT250206 Ceramium virgatum ME US KT250188 Ceramium botryocarpum IR MW354764 Ceramium virgatum NB CA KT250235 Ceramium pallidum NO MN184264 Ceramium pallidum NO MN184245 Ceramium siliquosum FR KJ960523 Ceramium pallidum NO MN184273 Ceramium sp. 3 NO MN184247 Ceramium diaphanum NC US KF367755 Ceramothamnion coulteri (M. coulteri) CA US KM254414 Ceramium ciliatum IRKJ 179929 Centroceras gasparrinii BR MW354765 Centroceras gasparrinii BR MN 105048 Centroceras gasparrinii BE KP2227 45 Centroceras clavulatum KO EU194971 Centroceras gasparrinii KO NC085551 Centroceras sp. 6 GB T RMK202411 Centroceras sp. 5 GB T R MK202400 Corallophila atlantica BR MW354774 Corallophila atlantica BR MW354775 Corallophila atlantica BR MW354761 Ceramieae sp. 5 BR MW354762 Corallophila eatoniana CA US HO544058 Gayliella jolyana BR MN105036 Gayliella sp. 9 HI US HO423004 Gayliella sp. 6 HI US HQ422894 Gayliella jolyana BR MN105033 Gayliella sp. 8 HI U S HO422973 Gayliella tamoiensis BR MN105028 Gayliella tamoiensis BR MN105025 Gayliella dawsonii BR MN105032 Gayliella ardissonei BR MN105046 Gayliella ardissonei BR MN105047 Gayliella sp. 5 BR MN105042 Corallophila eatoniana CA US KM254791 Gayliella sp. 4 BR MN105038 Gayliella iemanja BR MN1 05039 Centroceras sp. 7 HI US HO422953 Centroceras illaqueans BE KP222781 Centroceras hyalacanthum BE KP222789 Centroceras sp. 8 AU HM917544 Gayliella HO422977 Centroceras micracanthum BE KP2 22770 Centroceras arcii BE KP222761 Centroceras so. 9 CA KM254257 Pseudoceramium tenerrimum KO NC085558 Pseudoceramium inkyui KO NC085556 Pseudoceramium oliveirae BR MW354770 Campylaephora kondoi CH JO619158 Campylaephora kondoi KO NC085550 Ceramieae sp. 10 (Cer. diaphanum) CHKC795911 Campylaephora sungminbooi KO NC085557 Ceramieae sp. 9 (Campylaephora californica) CA US KM254681 Ceramieae sp. 2 (M. alternata) BG TR MK202416 Yoneshiguea affinis BR MW354769 Ceramothamnion sp. 2 BR MW354771 Ceramothamnion vagans BR MW354772 Ceramothamnion sp. KO OR8853 37 Ceramothamnion japonicum C H KC795912 Ceramium echionotum FR KJ960509 Ceramieae sp. 11 (Microcladia sp.) CA US KM254932 Ceramothamnion codii PBA1899 BE PP868169 Ceramothamnion fujianum BR MW3547 58 Ceramothamnion fujianum BR MW354759 Ceramothamnion brasiliense BR M W354760 Ceramieae sp. 8 (Cer. hanaense) HI US HO422844 Ceramieae sp. 7 (Cer. womersleyi) HI US HO423060 Carpoblepharis flaccida SA MH251749 Ceramothamnion dumosertum HI US HQ422603 Ceramium shuttleworthianum NO MN184486 Ceramieae sp. 4 BR MW354766 Ceramieae sp. 4 BR MW354767 Reinboldiella filicula BR MW354768 Ptilota serrata BC CA A Y970640 Aglaothamniom uruguayense BR MW354773 0.98/89 0.96/88 1.00/97 0.97/94 -/90 -/98 1.00/97 0.93/98 0.99/96 0.94/91 -/96 -/94 0.99/- 1.00/98 1.00/98 0.92/- 0.97/- 0.95/97 0.98/- 1.00/0.98 1.00/- 1.00/- 1.00/95 0.95/- 0.97/100 -/91 1.00/99 0.96/100 * * *** * * * * * * * * * * * * * 0.04 102 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Centroceras gasparrinii Centroceras clavulatum ARS03255 US: Hawaii — — — HQ422016 — Centroceras gasparrinii Centroceras clavulatum ARS03573 US: Hawaii — — — HQ422101 — Centroceras gasparrinii —TC826 USA: Texas, Port Aransas, Aransas Co 08.VII.2003 T.O. Cho & B.Y. Won —DQ374345 — Centroceras gasparrinii —TC367 USA: Florida, Sand Key State Park, St. Petersburg, Pinellas Co., W. Florida 19.IX.2002 T.O. Cho —DQ374346 — Centroceras gasparrinii —TC833 USA: Hawaii, Ka’ala’wai, O’ahu 08.VIII.2003 P. Vroom —DQ374354 — Centroceras gasparrinii — W00265 Panama: Isla Culebra, Pacific Panama 16.IV.1999 B. Wysor — DQ374359 — Centroceras gasparrinii (Meneghini) Kützing subsp. minor M.A. Wolf, A. Buosi, A.S. Juhmani & A. Sfriso Centroceras gasparrinii A000838 Italy: Venice — — LR536401 — — Centroceras hommersandii Won, T.O.Cho & Fredericq —B105 South Africa: Trafalgar, KwaZulu-Nata 20.II.1999 E. Coppejans et al. DQ374322 — — Centroceras hyalacanthum Kützing —TC524 USA: Florida, Nerritt Island, Brevard Co., E. Florid 05.III.2003 T.O. Cho & B.Y. Won DQ374341 DQ374366 — Centroceras hyalacanthum —BDA 0590 Bermuda: Cove east of Spanish Pt. 18.I.2012 C.W. Schneider et C.E Lane KP222803 —KP222789 Centroceras hyalacanthum —ARS02737 USA: Hawaii, Oahu — — — HQ422526 — Centroceras illaqueans C.W. Schneider, Cianciola & Popolizio —BDA 09-2-9 Bermuda: Capt. William´s Bay 15.III.2009 C.W. Schneider et C.E Lane KP222802 —KP222781 Centroceras micracanthum Kützing — W01063 Panama: Galeta Island, Smithsonian Tropical Research Institute (STRI), Caribbean entrance to the Panama Canal 01.X.1999 B. Wysor DQ374336 DQ374365 — Centroceras micracanthum Kützing —BDA 0584 Bermuda: Cove east of Spanish Pt. 18.I.2012 C.W. Schneider KP222804 —KP222770 Centroceras natalense Won, T.O. Cho, & Fredericq —TC2796 South Africa: Leisure Bay 24.VIII.2005 T.O. Cho & S. Fredericq DQ374343 — — Centroceras rodmanii Won, T.O. Cho & Fredericq —TC435 Chile: Cocholgue, Bahia Concepcion 01.I.1996 MHH DQ374333 DQ374364 — Centroceras tetrachotomum Won, T.O. Cho & Fredericq —TC415 South Africa: Lambert Bay, Western Cape 25.I.2001 MHH DQ374323 DQ374360 — Centroceras sp. 1 Centroceras clavulatum -Jamaica: Falmouth, Trelawny Province VII.1999 D.T. Thomas DQ022809 — — Appendix 1. — Continuation. 103 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Centroceras sp. 2 Centroceras sp. 1 HV06148 Australia: VIC, Mallacoota — Heroen Verbruggen, Kyatt Dixon MK125359 — — Centroceras sp. 3 Centroceras sp. 3 HV06015 Australia: VIC, Mallacoota — Heroen Verbruggen, Kyatt Dixon MK125362 Centroceras sp. 4 Centroceras clavulatum AC272 USA: CA, Monterey 07.IX.2006 -GQ252451 — — Centroceras sp. 5 Centroceras sp. 1 1Tris UK: Tristan da Cunha, Saint Helena: Nightingale. Nightingale 2010 024 21.III.2010 Sue Scott MK185797 —MK202400 Centroceras sp. 6 Centroceras sp. 1 1Tris UK: Tristan da Cunha, Saint Helena: West of Harbour, pools 2010 069 31.III.2010 Sue Scott — — MK202411 Centroceras sp. 7 —ARS03786 USA:Hawaii, Oahu 19.IV.2008 A. Kurihara — — HQ422953 Centroceras sp. 8 Centroceras clavulatum GWS015026 Australia 19.I.2010 G.W. Saunders & K. Dixon — — HM917544 Centroceras sp. 9 Centroceras clavulatum GWS021277 USA: California, Pigeon Point Lighthouse 15.V.2010 B. Clarkston & K. Hind — — KM254257 Ceramium barbatum Kützing —CAM 1528 Italy: Lerici, Liguria. BM000701867 11.VII.2001 CAM FR871396 — — Ceramium botryocarpum A.W.Griffiths ex Harvey —CAM 275 N. Ireland: Annalong, Co. Down XI.1993 CAM AF439288 — — Ceramium botryocarpum Roth —CAM 1081 Annalong, Co. Down, N. Ireland 25.I.2003 CAM — MW354748MW354764 Ceramium botryocarpum Roth —CAM 1086 Annalong, Co. Down, N. Ireland 25.I.2003 Ellen Cocquyt —PQ047457 — Ceramium ciliatum (Ellis) Ducluzeau — GALW Ireland: Finavarra, Co. Clare 2003 L.M. GALW FR871397 — — Ceramium ciliatum (Ellis) Ducluzeau —strain 25996 Spain: Tragove, Pontevedra 04.III.2011 Javier Cremades — — KJ179929 Ceramium cimbricum H.E.Petersen in Rosenvinge —CAM 211 Ireland: Tamney, Mulroy Bay, Donegal. BM000701865 (Maggs & Hommersand, 1993, Fig 15b) 15.II.1993 CAM MW366531 — — Ceramium cimbricum H.E.Petersen in Rosenvinge —CAM 212 Ireland: Tamney, Mulroy Bay, Donegal. BM000701863 15.II.1993 CAM MW366532 — — Ceramium cimbricum H.E.Petersen in Rosenvinge —C 8843/C-A37605 Denmark: Egerslev Ron, Limfjorden 27.VII.1920 H. Petersen KU925350 — — Ceramium circinatum (Kützing) J.Agardh —CAM 181 England: Kimmeridge, Dorset 13.X.1992 CAM FR871398 — — Ceramium connivens Zanardini —-Italy: Ca’ Roman, Pellestrina —-LR031254 — — Ceramium derbesii Solier ex Kützing — #4 (HA 3150-1) Venice Lagoon, Lido di Veneza, Italy 00.VI.2010 Wolf & Sciuto FR775778 — — Ceramium deslongchampsii Chauvin ex Duby —CAM 45 N. Ireland: Helen’s Bay, Co. Down 30.III.1992 CAM AF439293 — — Appendix 1. — Continuation. 104 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Ceramium diaphanum (Lightfoot) Roth —NCweed1220 USA: North Carolina, Onslow Bay, Atlantic Ocean, Wynne Plaza Park 18.III.2013 D.W. Freshwater — — KF367755 Ceramium diaphanum sensu Harvey —CAM 199 England: Cornwall, 10.III.1993 CAM FR871401 — — Ceramium aff. diaphanum —CAM 249 Canada: Pomquet Harbour, Antigonish Co., Nova Scotia 26.IX.1993 C. Bird FR871400 — — Ceramium aff. diaphanum —CAM 136 Ireland: St John’s Point, West Donegal 29.VIII.1992 CAM FR871399 — — Ceramium echionotum J.Agardh —LM 1027 Canary Islands: Corallejo, Fuertaventura 25.III.1999 CAM AF439313 — — Ceramium echionotum J.Agardh —RMAR2945 France: Brittany, Le Loup 30.VIII.2011 Couceiro L., Robuchon M. — — KJ960509 Ceramium gaditanum (Clemente) Cremades —CAM 1052 UK: Broad Haven, Pembrokeshire, Wales. BM013821504 18.III.2000 CAM MW366534 — — Ceramium gaditanum —MBBB 64 Spain: Cudillero, Asturias. 20.I.2005 CAM — MW354753 — Ceramium glandulosum (Solander ex Turner) Barros-Barreto & Maggs Microcladia glandulosa (Solander ex Turner) Greville CAM 322 England: Helston, Cornwall BM(NH) 30.VII.1994 CAM MW366533 — — Ceramium nodosum (Kützing) A.W. Griffiths & Harvey — — Italy: Ca’ Roman, Pellestrina — — LR031256 — — Ceramium nudiusculum (Kützing) Rabenhorst — #2 (HA 3150-4) Venice Lagoon, Lido di Veneza, Italy 00.VI.2010 Wolf & Sciuto FR775780 — — Ceramium pallidum (Kützing) Maggs & Hommersand —CAM 223 Ireland: Flannery’s Bridge, Kilkieran Bay, Co. Galway 12.VI.1993 CAM AF439292 — — Ceramium pallidum —CAM 1071 UK: Northern Ireland, Strangford Lough, The Dorn. 24.X.2002 Ellen Cocquyt —PQ047456 — Ceramium pallidum — GWS039884 Norway: Ormhilleren (site 1) 05.VI.2016 G.W. Saunders & T. Bringloe MN184533 — MN184264 Ceramium pallidum GWS039872 Norway: Hellesoy 05.VI.2016 G.W. Saunders & T. Bringloe — — MN184245 Ceramium pallidum 2016_ BIO309A_47 Norway: Hordaland, Sore egdholmen 18.IV.2016 C.M. Berntsen, N. E. Frogg, C.C. Martin — — MN184273 Ceramium rothianum M.A.Wolf, K.Sciuto, I. Moro, Maggs & Sfriso — — Italy: Ca’ Roman, Pellestrina — — LR031258 — — Ceramium secundatum Lyngbye —CAM 41 Ireland: St John’s Pt, Co. Donegal 20.II.1992 CAM AF439287 — — Ceramium secundatum —CAM 355 Dale Fort, Pembs, Wales 10.X.1995 CAM AF439286 — — Appendix 1. — Continuation. 105 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Ceramium secundatum — — France: Bretagne, Roscoff 05.IV.2000 Yang,E.C. Boo,S.M. — — EU194972 Ceramium secundatum —IN07.R036 UK:Tristan da Cunha. Saint Helena, Inaccessible, Off Tom`s Beach 29.XI.2007 Susan Scott MK185826 — — Ceramium secundatum —Tristan 2010 073 Saint Helena: Tristan, West of Harbour, pools 31.III.2010 Susan Scott — — MK202399 Ceramium secundatum —Nightingale 2010 056 Saint Helena: Nightingale, East Landing 23.III.2010 Susan Scott — — MK202407 Ceramium secundatum — GWS018041 US: Fort Witherill, Jamestown, Rhode Island 16.IV.2016 B. Clarkston, D. McDevit, M. Bruce, A. Savoie, C. Longtin KT250273 —KT250208 Ceramium secundatum —2016_ BIO309A_30 Norway: Hordaland, Espegrend 17.IV.2016 C.S. Armitage — — MN184367 Ceramium secundatum — GWS038089 Norway: Station Dorm Beach, Espegend 03.VI.2016 G.W. Saunders & T. Bringloe — — MN184499 Ceramium shuttleworthianum (Kützing) Rabenhorst —CAM 1049 UK: Broad Haven Beach, Wales 17.III.2003 CAM FR871404 — — Ceramium shuttleworthianum — GWS039860 Norway: Hellesoy 05.VI.2016 G.W. Saunders & T. Bringloe MN184563 — MN184486 Ceramium siliquosum (Kutzing) Maggs & Hommersand —CAM 172 Ireland: Fanore, Co. Clare 28.IX.1992 CAM FR871405 — — Ceramium siliquosum —RMAR1120 France: Brittany, Les Rospects 14.III.2011 Le Gall, L., Turpin, Y. — — KJ960523 Ceramium strictum Roth —CAM 140 Ireland: Skerries, Co. Dublin 30.VIII.1992 CAM FR871408 — — Ceramium tenuicorne (Kützing) Waern Rueness 9817 Culture 04.X.1994 B. Eklund FR871409 — — Ceramium virgatum Roth —CAM 496 Germany: South Harbour, Helgoland, North Sea 07.I.1999 A. Wagner AF439290 — — Ceramium virgatum — GWS003615 USA: Maine, Cape Neddick, southern ME 24.IV.2006 L. Le Gall — — KT250188 Ceramium virgatum — GWS014784 USA: Massachusetts, Folly Cove, Gloucester, right side 13.IV.2010 B. Clarkston, A. Savoie KT250272 —KT250206 Ceramium virgatum — GWS002366 or GWS2366 Canada: Nova Scotia, Cape St. Marys 5.IX.2004 G.W. Saunders —EF033604 KT250235 Ceramium sp. 1 Ceramium sp. 2016_ BIO309A_66 Norway: Hordaland, Stora Kalsoy 19.IV.2016 T. Ulvatn, H. Dybevik MN184543 — MN184330 Ceramium sp. 2 Ceramium secundatum LM 1565 New Zealand: Christchurch, Cultured I.2003 M.P. Johnson MW366509 MW354749MW354763 Ceramium sp. 3 Ceramium sp1 GWS040811 Norway: Hakonsund 10.VI.2016 G.W. Saunders & T. Bringloe MN184529 — MN184247 Ceramium sp. 4 —LM 1562 Tenerife, Spain 2003 LM MW366507 — — Appendix 1. — Continuation. 106 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Ceramium sp. 5 —CAM 1130 Italy: Ischia. Culture 7.XII.2002 CAM FR871407 — — Ceramothamnion brasiliensis (A. B. Joly) M.J.Wynne & C.W.Schneider Ceramium brasiliense A.B. Joly MBBB 41 Brazil: Boa Viagem, Recife, Pernambuco. RB420020 15.XII.2001 J. Torres & L. Gestinari DQ458922 MW354747MW354760 Ceramothamnion codii H.Richards — — Bermuda: Cooper´s Island P.B.-A. 1899 from BM000936098 29.IV.1912 F.S. Collins PP868167 PP863326 PP868169 Ceramothamnion codicola (J.Agardh) M.J.Wynne & C.W.Schneider Ceramium codicola J.Agardh —USA: Boiler Bay, Oregon 08.VII.1998 T.O. Cho & G.I. Hansen AY155522 DQ238806 — Ceramothamnion coulteri (Harvey) M.J.Wynne & C.W.Schneider Microcladia coulteri Harvey CAM 402 USA: Asilomar Beach, California. BM013821501 00.VII.1996 CAM MW366528 — — Ceramothamnion coulteri Microcladia coulteri Harvey AC263 USA: WA, San Juan Is. 18.VIII.2006 Carlile, A. L. GQ252482 — — Ceramothamnion coulteri Microcladia coulteri Harvey GWS022133 USA: California, Stillwater Cove, Pebble Beach 20.V.2010 B. Clarkston, K. Hind & S. Toews — — KM254414 Ceramothamnion dumosertum (R.E. Norris & I.A. Abbott) M.J.Wynne & C.W.Schneider Ceramium dumosertum R.E. Norris & I.A. Abbott AC191 USA: Hawaii,Kihei, Maui 05.IV.2006 Carlile, A. L. GQ252454 — — Ceramothamnion dumosertum Cer. dumosertum R.E. Norris & I.A. Abbott ARS00847 USA: Hawaii, Maui, Lipoa St 05.IV.2006 Carlile, A. L. —HQ421685HQ422603 Ceramothamnion fujianum (BarrosBarreto & Maggs) M.J.Wynne & C.W.Schneider Ceramium fujianum BarrosBarreto & Maggs MBBB 22 Brazil: Itacuruça, Cananéia, Sao Paulo. RB420061 16.VI.2003 M.T. Fujii DQ458949 MW354750MW354758 Ceramothamnion fujianum Ceramium fujianum MBBB 82 Brazil: Praia Baleia, Manguinhos, Espirito Santo. RB420063 26.III.2005 M.T. Fujii — — MW354759 Ceramothamnion fujianum Ceramium fujianum MBBB 84 Brazil: Portal de Santa Cruz, Espirito Santo. RB420064 25.III.2005 M.T. Fujii DQ458952 — — Ceramothamnion horridulum (P.C.Silva) M.J.Wynne & C.W.Schneider Ceramium horridum Setchell & N.L. Gardner — México: San Juan de la Costa, Baja Califórnia 15.VI.2000 T.O. Cho & R.R.- Rodriguez AF521796 DQ238807 — Ceramothamnion inconspicuum (Zanardini) M.J.Wynne & C.W.Schneider Ceramium incospicuum Zanardini —Alberoni dyke, Lido of Venice — — LR031255 — — Ceramothamnion interruptum (BarrosBarreto & Maggs) M.J.Wynne & C.W.Schneider Ceramium interruptum Setchell & Gardner —USA: Dana Point, California 04.XII.1999 T.O. Cho & S. Murray; AY155527 — — Ceramothamnion japonicum (Okamura) M.J.Wynne & C.W.Schneider Ceramium japonicum Okamura —South Korea: Namhae, Kyengnam 03.III.1998 T.O. Cho AY948753 — — Appendix 1. — Continuation. 107 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Ceramothamnion japonicum Ceramium japonicum Okamura LQDHQW20121115R5 China: Huiquan Bay, Qingdao 11.XI.2012 Wu F.-F. — KC795870 — Ceramothamnion japonicum Ceramium japonicum Okamura CQDHQW20121115R5 China: Huiquan Bay, Qingdao 21.XI.2012 Wu F.-F. — — KC795912 C. julieae (A.Millar & P.G.Richards) M.J.Wynne & C.W.Schneider Ceramium juliae A.J.K. Millar — South Africa culture W. Stirk MW366506 — — Ceramothamnion pacificum (Collins) M.J.Wynne & C.W.Schneider Ceramium pacificum (Collins) Kylin AC306 USA: Washington, San Juan Island 29.VII.2007 Carlile, A. L. FJ795539 — — Ceramothamnion pacificum Ceramium pacificum (Collins) Kylin AC286 USA: WA, Clallam Co., Cape Alava 18.VI.2007 Carlile, A. L. GQ179817 — — Ceramothamnion riosmenae (B.Y.Won & T.O.Cho) M.J.Wynne & C.W.Schneider Ceramium riosmenae B.Y.Won & T.O.Cho TC245 Mexico: B.C.S., Bahía Magdalena, López Nateos, Estero Santo Domingo 01.VI.2000 T.O.Cho & R.R.- Rodriguez JN712693 — — Ceramothamnion sinicola (Setchell & N.L.Gardner) M.J.Wynne & C.W.Schneider Ceramium sinicola Setchell & Gardner DANA USA: Dana Point, California 04.XII.1999 T.O. Cho & S. Murray; AY155531 — — Ceramothamnion sinicola Ceramium sinicola AC298 USA: CA, Santa Cruz Co., Greyhound Rock 03.VII.2007 —GQ179818 — — Ceramothamnion vagans (P.C.Silva) M.J.Wynne & C.W.Schneider Ceramium vagans P.C. Silva MBBB 126 Brazil: Praia da Feiticeira, Ilha Grande, Mangaratiba, Rio de Janeiro. RB420069 29.VII.2005 MBBB MW366529 — MW354772 Ceramothamnion sp. 1 —CAM 1144 Caldera, Chile. BM013821505 undated Juan Correa MW366508 — — Ceramothamnion sp. 2 —MBBB 110 Brazil: Prainha, Arraial do Cabo, Rio de Janeiro. RB420052 27.VII.2005 MBBB MW366530 — MW354771 Ceramothamnion sp. 3 Ceramium sp. 2 DWF-2013 USA: North Carolina, Onslow Bay, Atlantic Ocean, Wynne Plaza Park NCweed1227 18.II.2013 D.W. Freshwater KF367768 — — Ceramothamnion sp. 4 Ceramium sp. ARS01431 USA: Hawaii — — — HQ422441 — Ceramothamnion sp. 5 Ceramium sp. ARS03593 USA: Hawaii — — — HQ422096 — Ceramothamnion sp. 6 Ceramium codii (H.Richards) Mazoyer — USA: Hawaii, Kure Atoll 08.VII.1998 T.O. Cho & G.I. Hansen —HQ421967 — Corallophila atlantica (A.B.Joly & Ugadim) R.E.Norris —MBBB 146 Brazil: Prainha, Arraial do Cabo, Rio de Janeiro. RB420071 27.VII.2005 MBBB MW366512 — MW354761 Appendix 1. — Continuation. 108 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Corallophila atlantica —MBBB 457 Brazil: Praia da Tartaruga, Armação de Búzios, Rio de Janiero 11.IX.2015 MBBB — — MW354774 Corallophila atlantica MBBB 458 Brazil: Praia da Tartaruga, Armação de Búzios, Rio de Janiero 11.IX.2015 MBBB MW366536 — MW354775 Corallophila eatoniana (Farlow) T.O.Cho, H.-G. Choi, G.I.Hansen & S.M.Boo Ceramium eatonianum (Farlow) De Toni GWS021384 USA: California, Pigeon Point Lighthouse 15.V.2010 B. Clarkston & K. Hind — — KM254535 Corallophila eatoniana —XX1109 California, USA — E. Serrão DQ458957 — — Corallophila eatoniana —TC047 USA: Friday harbor, Washington, 16.VII.1998 T.O. Cho AY945765 DQ238802 — Corallophila eatoniana —AC135 Canada: B.C., Vancouver Is. 05.VII.2004 AC GQ252469 — — Corallophila eatoniana — — USA: Oregon, Crystal Core 15.XII.1999 —DQ787568 — — Corallophila eatoniana Ceramium eatonianum GWS021391 USA: California, Santa Cruz (Four Mile) 15.V.2010 B. Clarkston, & K. Hind — — HQ544058 Corallophila eatoniana Ceramium eatonianum GWS021384 USA: California, Pigeon Point Lighthouse 15.V.2010 B. Clarkston, & K. Hind — — KM254535 Corallophila huysmansii (Webervan Bosse) R.E.Norris —ARS03236 USA: Hawaii, Kure Atoll 30.XI.1999 I.A. Abbott Collection —HQ421935HQ422782 Corallophila sp. Ceramium sp. ARS03774 USA: Hawaii — — — HQ422146 — Gayliella ardissonei Barros-Barreto & F.P.Gomes Ceramium flaccidum MBBB 26 Brazil: Ponta da Baleia, Aracruz, Espirito Santo. RB420056 07.VII.2002 MBBB DQ458944 — — Gayliella ardissonei Ceramium flaccidum MBBB 50 Brazil: Praia Mole, Vitória, Espirito Santo. RB420057 01.VIII.2003 MBBB — — MN105046 Gayliella ardissonei Ceramium flaccidum MBBB 51 Brazil: Praia Mole, Vitória, Espirito Santo. RB420058 01.VIII.2003 MBBB DQ458946 MN105050MN105047 Gayliella dawsonii (A.B.Joly) BarrosBarreto & F.P.Gomes Ceramium dawsonii MBBB 53 Brazil: Redonda Sul, Abrolhos, Bahia. RB420047 14.XII.2002 MBBB DQ458938 — MN105032 Gayliella dawsonii —FPG234 Praia do Gonzaguinha, São Vicente, São Paulo. Brazil. R232583 12.X.2016 F. P. Gomes — MN105057 Gayliella fimbriata (Setchell & N.L.Gardner) T.O.Cho & S.M.Boo —AC202 USA: HI, Maui 08.IV.2006 —GQ252474 — — Gayliella flaccida (Harvey ex Kützing) T.O.Cho & L.J.McIvor —CAM 113 UK: Beau Port, Jersey, Channel Islands 12.VIII.1992 CAM AY945778 — — Gayliella iemanja Barros-Barreto & F.P.Gomes Ceramium dawsonii MBBB 34 Brazil: Serrambi, Pernambuco. RB420045 14.XII.2001 J. Torres & L. Gestinari DQ458936 MN105052MN105039 Appendix 1. — Continuation. 109 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Gayliella jolyana Barros-Barreto & F.P.Gomes Ceramium dawsonii MBBB 49 Brazil: Peracanga, Guarapari, Espirito Santo. RB420046 31.VII.2003 MBBB DQ458937 MN105061MN105033 Gayliella jolyana Ceramium dawsonii MBBB 69 Brazil: Prainha, Arraial do Cabo, Rio de Janeiro. RB420050 29.VII.2001 MBBB — MN105060MN105036 Gayliella mazoyerae T.O. Cho, Fredericq & Hommersand — — Italy: Sicily XII.2003 Cormaci AY945776 DQ238809 — Gayliella nanaoensis Gayliella sp. HTa—2020a China — Tan Huaqiang MN592682 — — Gayliella tamoiensis Barros-Barreto & F.P.Gomes —MBB 241 Brazil: Praia Rasa, Armacao dos Buzios, RJ R232576 2.VI.2011 MBBB MN105075 MN105055MN105028 Gayliella tamoiensis —MBB 434 Brazil: Praia Rasa, Armacao dos Buzios, RJ R232578 2.VI.2011 MBBB — MN105056MN105025 Gayliella taylorii (E.Y. Dawson) T.O. Cho & S. M. Boo —TC255 USA: Crystal Cove, Orange Co., California 3.VII.1999 T.O. Cho & S.N. Murray AY945774 — — Gayliella taylorii (E.Y. Dawson) T.O. Cho & S. M. Boo — — USA: Laguna Beach, Orange Co., California 3.XII.1999 T.O. Cho & S.N. Murray —DQ238812 — Gayliella transversalis (Collins & Hervey) T.O. Cho & Fredericq Gayliella keywestiensis TC961 USA: Key West, Florida 30.XI.2003 T.O. Cho AY945777 DQ238810 — Gayliella womersleyi T.O. Cho, Maggs & L. McIvor Gayliella miniatum C1507 Australia: Williamstown 13.III.2002 M.D. Guiry AY945780 DQ238808 — Gayliella sp. 1 Gayliella sp. AC165 Mexico: Baja CA, Cabo San Quintin 20.III.2005 GQ252475 Gayliella sp. 2 — JAW 4461 Williamstown, South Australia, culture 24.VII.2004 J.A.West MW366510 — — Gayliella sp. 3 Gayliella sp. 1 JFC—2019 HV06133 Australia: VIC, Mallacoota — Heroen Verbruggen, Kyatt Dixon MK125376 — — Gayliella sp. 4 —FPG 202 Brazil: Praia do Gonzaguinha, Sao Vicente, SP R232573 12.X.2016 F.P. Gomes MN105065 MN105054MN105038 Gayliella sp. 5 —FPG 125 Brazil: Praia da Pedra Rachada, Paracuru, CE R232568 4.VI.2016 F.P. Gomes — MN105051MN105042 Gayliella sp. 6 —ARS03583 USA: Hawaii, Molokai, Wavecrest 21.III.2008 K. Conklin —HQ422107HQ422894 Gayliella sp. 7 —ARS03693 USA: Hawaii, Lanai 28.III.2008 A. Kurihara —HQ422193 HQ422977 Gayliella sp. 8 —ARS03689 USA, Hawaii, Oahu, Nanakuli 12.III.2008 A. Kurihara —HQ422171HQ422973 Gayliella sp. 9 —ARS03846 USA, Hawaii, Oahu, Makai Pier 08.VI.2008 A. Kurihara —HQ422195HQ423004 Herpochondria borealis (Ruprecht) Barros-Barreto & Maggs Microcladia borealis Ruprecht CAM 424 USA: Salt Creek, Crescent Bay, Washington NCU-A-0020296 09.VI.1994 MHH MW366515 — — Appendix 1. — Continuation. 110 CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Barros-Barreto B. et al. Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Herpochondria borealis Microcladia borealis TC391 USA: Lone Ranch Beach, Brookings, Josephine Co, Oregon 18.V.1999 MHH EF613514 — — Herpochondria borealis Microcladia borealis —USA: Otter Crest, Oregon 12.VII.1998 T.O. Cho & G.I. Hansen —DQ238799 — Herpochondria corallinae (G. Martens) Falkenberg — H6 South Korea: Jeongdori, Wando 10.VII.1998 S.M. Boo, T.O. Cho & H.G. Choi DQ787569 — — Herpochondria elegans (Okamura) Itono —TC037 South Korea: Jeongdori, Wando 15.XII.1998 S.M. Boo, T.O. Cho & H.G. Choi AY945771 DQ238800 — Pseudoceramium brevizonatum (H.E. Petersen) BarrosBarreto & Maggs —CAM 240 Italy: Pietro Ligure, Liguria. BM000701867 19.VII.1993 CAM MW366524 — — Pseudoceramium caraibicum (H. E. Petersen & Borgesen) BarrosBarreto Maggs & M.A.Jaram. Ceramium brevizonatum H. E. Petersen — Mexico: Yucatan 13.II.1993 C.F.D. Gurgel AF259491 — — Pseudoceramium clarionense (Setchell & N.L. Gardner) BarrosBarreto & Maggs Ceramium clarionense Setchell & N.L. Gardner MBBB 76 Brazil: Siriba, Abrolhos, Bahia. RB420036 13.XII.2002 MBBB DQ458933 Pseudoceramium inkyui (T.O. Cho, Fredericq & S.M. Boo) BarrosBarreto & Maggs Ceramium inkyui T.O. Cho, Fredericq & S.M. Boo —South Korea: Sinnam, Kangwon 7.VII.1999 T.O.Cho MW366527 — — Pseudoceramium paniculatum (Okamura) BarrosBarreto & Maggs Ceramium paniculatum Okamura —South Korea: Gampo, Kyungbuk 8.IX.1999 S.M. Boo & T.C. Cho AF521802 — — Pseudoceramium planum (Kützing) Barros-Barreto & Maggs Ceramium planum Kützing CAM 413 South Africa: Kommetjie, Western Cape 9.XI.1999 MHH MW366527 — — Pseudoceramium oliveirarume Barros-Barreto, M.A.Jaram. & Maggs Ceramium brevizonatum var. caraibicum H.E. Petersen & Børgesen MBBB 47 Brazil: Costão, Itapemirim, Espirito Santo. RB420029 04.III.2004 MBBB — MW354756 — Pseudoceramium oliveirarume Ceramium brevizonatum v. caraibicum MBBB 98 Brazil: Ponta do Pái Vitório, Armação dos Búzios, Rio de Janeiro. RB420031 10.VII.2005 MBBB — — MW354770 Pseudoceramium oliveirarume Ceramium brevizonatum v. caraibicum MBBB 80 Brazil: Guaibura, Guarapari, Espirito Santo. RB420030 31.VII.2003 MBBB DQ458932 — — Pseudoceramium tenerrimum (G. Martens) BarrosBarreto & Maggs Ceramium tenerrimum —South Korea: Namhae, Kyungnam 30.V.1999 T.O.Cho & H.S.Yoon AF521805 — — Pseudoceramium tenerrimum Ceramium tenerrimum AC209South Korea: Jeju City, Yongdam 11.V.2006 —GQ252467 — — Pseudoceramium tenerrimum Ceramium tenerrimum —South Korea: Gampo, Kyengju 08.IX.1999 T.O. Cho —DQ238805 — Appendix 1. — Continuation. 111 Synonymy of Stirkia Barros-Barreto & Maggs with Ceramothamnion H.Richards (Rhodophyta) CRYPTOGAMIE, ALGOLOGIE • 2025 • 46 (7) Sample Herbaria/ GenBank names Sample number Location and specimen voucher Date collected Collector GenBank accession number rbcLPartial LSU COI-5P Pseudoceramium tenerrimum Ceramium tenerrimum LQDHQW20120309R14 China: Huiquan Bay, Qingdao 12.V.2012 Wu F.-F. — KC795867 — Pseudoceramium sp. 1 —CAM 1654 Marigot, St Martin, Dutch Antilles. BM013821502 19.I.2005 W. Prud’homme van Reine MW366523 — — Pseudoceramium sp. 2 Ceramium tenerrimum MBBB 77 Redonda Sul, Abrolhos, BA; Brazil. RB420068 14.XII.2002 MBBB DQ458954 — — Reiboldiella filicula (Harvey ex Womersley) BarrosBarreto & Maggs Ceramium filicula Harvey ex Womersley XX1015 Australia: Melbourne, Williamstown — J.C. Zuccarello DQ458956 — — Reinboldiella filicula Ceramium filicula MBBB 17 Brazil: Praia Vermelha, Urca, Rio de Janeiro. RB420054 13.IX.2004 W. Paradas DQ458942 — MW354768 Reinboldiella orientalis S.-M.Lin & Mas. Suzuki — RO-CJ3 Taiwan: Keelung, Chaojing 05.V.2016 S.-M. Lin LC201458 — — Reinboldiella orientalis — RO-CJ4 Taiwan: Keelung, Chaojing 05.V.2016 S.-M. Lin LC201459 — — Reinboldiella schmitziana (Reinbold) De Toni —TC032 South Korea: Guryongpo, Pohang 04.V.1998 T.O.Cho & H.S. Yoon AY945772 — — Reinboldiella schmitziana — RS-JP South Korea: Guryongpo, Pohang 05.VI.2009 S.-M. Lin & M. Suzuki LC201465 — — Reinboldiella taiwanensis S.-M. Lin & Mas. Suzuki — RT-CJ1 Taiwan: Chaojing, Keelung 07.IV.2016 S.-M. Lin LC201467 — — Reinboldiella taiwanensis — RT-CJ2 Taiwan: Chaojing, Keelung 07.IV.2016 S.-M. Lin LC201468 Reinboldiella warburgii (Heydrich) Yoshida & Mikami — — Taiwan: St. Tiago, Taipei Co. 22.V.2005 S.-M. Lin EF125878 — — Reinboldiella warburgii — RW-CJ Taiwan: Chaojing, Keelung 28.IV.2009 M. Suzuki LC201471 — — Yoneshiguea affinis (Setchell & N.L. Gardner) BarrosBarreto & Maggs Ceramium affine Setchell & N.L. Gardner —México: Pichilingue, Baja Califórnia, 27.V.2000 T.O. Cho & R.R. Rodriguez AF521797 DQ238803 — Yoneshiguea affinis (Setchell & N.L. Gardner) BarrosBarreto & Maggs Ceramium affine MBBB 65 Brazil: Lagoinha,Ubatuba, Sao Paulo. RB420018 24.V.2003 M.T. Fujii DQ458920 MW354755MW354769 Yoneshiguea compta (Børgesen) BarrosBarreto & Maggs Ceramium comptum Børgesen MBBB 118 Brazil: Lagoa Azul, Ilha Grande, Mangaratiba, Rio de Janeiro. RB420041 29.VII.05 MBBB MW366522 — — Yoneshiguea sp. — JAW 3507 French Polynesia: Cooks Bay, Moorea, Culture 01.VII.1995 J.A. West MW366521 — — Ceramieae sp. 1 —CAM 1133 France: Thau Lagoon VI.2005 M. Verlaque MW366525 — — Ceramieae sp. 2 Microcladia alternata Nightingale 2010 040 UK: Tristan da Cunha, Saint Helena: Nightingale, Southwest Point 22.III.2010 Susan Scott MK185813 —MK202416 Ceramieae sp. 3 Ceramium tenuissimum (Roth) J.Agardh AC213 South Korea: Jeju Is., Onpyong 12.V.2006 —GQ252468 — — Appendix 1. — Continuation.