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Article

Integrative Approach to Species Delimitation in Sargassum (Fucales, Phaeophyceae) from Central American Pacific Based on Morphological and Genetic Evidence

by
Mariana Viales-Cubillo
1,2,3,*,
Fabio Quesada-Perez
1,2,
Paola Díaz-Canales
1,2,
Kaylen González-Sánchez
1,2 and
Cindy Fernández-García
1,2,4
1
Centro de Investigaciones en Ciencias del Mar y Limnología (CIMAR), Universidad de Costa Rica, San Pedro, San José 10905, Costa Rica
2
Posgrado de Biología, Sistema de Estudios de Posgrado, Universidad de Costa Rica, San José 10905, Costa Rica
3
Escuela de Biología, Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa Rica, San José 10905, Costa Rica
4
Posgrado en Gestión Integrada de Áreas Tropicales, Sistema de Estudios de Posgrado, Universidad de Costa Rica, San José 10905, Costa Rica
*
Author to whom correspondence should be addressed.
Diversity 2025, 17(9), 592; https://doi.org/10.3390/d17090592 (registering DOI)
Submission received: 3 July 2025 / Revised: 28 July 2025 / Accepted: 29 July 2025 / Published: 23 August 2025
(This article belongs to the Special Issue Eco-Physiology of Shallow Benthic Communities)
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Abstract

The genus Sargassum is taxonomically complex and poorly studied in the Eastern Tropical Pacific. We analyzed specimens collected along the Pacific coast of Central America and compared them with historical records and herbarium material to clarify species identities. Using detailed morphological analyses with molecular phylogenetic reconstruction based on concatenated ITS2 (Internal Transcribed Spacer 2) and COX3 (Cytochrome Oxidase Subunit 3) sequences, we identify two distinct morphotypes corresponding to two well-supported clades. One clade matches the morphology and molecular profile of Sargassum liebmannii. We provide the most comprehensive description of this species to date, including the first published ITS2 and COX3 sequences. Since Taylor’s 1945 work on the tropical Pacific of the Americas, S. liebmannii has been widely reported and considered the predominant species. It forms a genetic clade with other species from the Gulf of California; therefore, we propose a new section, Herporhizum/Sinicola. The second clade represents a previously unrecognized taxon from Central America, which we describe as a new species: Sargassum lacrucense, within the subgenus Sargassum, section Sargassum. Contrary to previous reports, Sargassum brandegeei—now recognized as Sargassum herporhizum—was not found in the region. This study underscores the importance of integrating morphological and molecular data to resolve Sargassum taxonomy in Central America.

1. Introduction

Brown algae (Phaeophyceae, Heterokontophyta) of the order Fucales are widely distributed in coastal areas around the world, with particularly high diversity in tropical and intertropical regions. In these environments, they form underwater forests that constitute structurally complex ecosystems of great ecological importance [1,2,3,4,5]. Their taxonomic classification is mainly based on morphological characteristics and their haplontic life cycles; however, this is complicated by the marked phenotypic plasticity and high level of polymorphism exhibited by many of their species [6].
Within the order Fucales, the genus Sargassum is the most diverse and is known for its high morphological variability, which has led to numerous taxonomic challenges and ambiguities in species delimitation [7,8,9,10]. This difficulty in identification can have both ecological and biotechnological implications, as some species contain bioactive compounds of interest, and misidentification can lead to errors in their selection [6].
Given these limitations, several studies have promoted the use of molecular tools, such as DNA (deoxyribonucleic acid) barcoding, to support taxonomic identification and phylogenetic studies within the genus Sargassum [8,11]. This technique has proven useful due to the small amount of material required and its high resolution at different taxonomic levels [12,13].
In the Pacific of Central America, along the coasts of Costa Rica and Nicaragua, seasonal Sargassum forests (Figure 1) have been recorded, with significant coverage during the colder months of the year (November to February), associated with upwelling events that increase nutrient availability [14,15,16]. These macroalgal forests play a crucial ecological role, serving as feeding grounds and nursery habitats for a wide variety of invertebrates, including mollusks, crustaceans, and polychaetes. The structural complexity of Sargassum provides shelter and food resources that support rich and diverse benthic communities. In Costa Rica, Sargassum ecosystems have been recognized as conservation targets in marine spatial planning and ecosystem-based management analyses conducted by the Ministry of Environment due to their ecological importance and vulnerability to environmental change [17].
Regarding specific composition, Sargassum liebmannii J. Agardh has been reported along the coasts of Costa Rica, Panama, Nicaragua, and El Salvador [14,18,19,20,21,22]. Beyond Central America, the species has also been recorded in North America, including Baja California, the Gulf of California, the Islas Revillagigedo, other regions of the tropical Pacific coast of Mexico [1,20,23,24,25,26], and in South America, specifically in Brazil [27] and Colombia [1]. The type locality of Sargassum liebmannii is San Agustín, Mexico. The isotype specimen (Figure A1) is incomplete, lacking a holdfast, but includes reproductive structures. Additionally, Sargassum brandegeei Setchell and N.L.Gardner has been documented in Costa Rica [20] and Sargassum sinicola Setchell and N.L.Gardner in Nicaragua [21].
Recent observations suggest the existence of a single dominant morphotype in some of these locations [28], highlighting the need for more detailed studies. Since the observed morphological variation could reflect unrecognized interspecific differences, it is essential to integrate morphological and molecular tools to clarify the species diversity of the genus in this region. In this study, we assessed the morphological variation and genetic structure of Sargassum populations collected in the northern Pacific of Costa Rica, complemented with specimens from Panama, Nicaragua, and El Salvador. To this end, we used two molecular markers (COX3 and ITS2) and a set of morphological measurements to explore species delimitation in this region of the Central American Pacific. Specifically, we aim to (i) clarify the taxonomy of the collected Sargassum specimens based on phylogenetic and morphological analyses and (ii) provide a reference framework for the identification of Sargassum species in the region.

2. Materials and Methods

2.1. Study Area and Sample Collection

Specimens of the genus Sargassum were collected along the northern Pacific coast of Costa Rica during April and June of 2023, 2024, and 2025 (Figure 2), in localities characterized by the presence of seasonal macroalgal forests. All collected specimens corresponded to mature individuals, most of them in reproductive condition. Collection was carried out by snorkeling, SCUBA(Self-Contained Underwater Breathing Apparatus) diving, or manual gathering, depending on the depth at which the specimens were found. At each site, morphotypes were preliminarily identified in situ based on observable morphological traits, following previously published taxonomic descriptions. In addition, collections from Costa Rica, Nicaragua, Panama, and El Salvador form the collection of the Phycology Laboratory at the University of Costa Rica (Figure 2), as well as specimens from various herbaria (UC, MICH, USJ), were used.
For molecular analyses, between 100 and 150 mg of tissue per sample was collected and stored frozen until laboratory processing. In addition, one well-preserved specimen of each morphotype, along with a duplicate, was selected for inclusion in the institutional herbarium. Complete specimens were photographed and labeled for later morphological analysis.
All collected specimens were deposited in the USJ Herbarium at CIBET, University of Costa Rica.

2.2. Morphological Characterization

A detailed morphological analysis was conducted using at least 10 representative specimens of each collected morphotype. For each specimen, the total thallus length was measured, and ten leaves were selected from each thallus. For these leaves, length and width were recorded, along with the number and arrangement of cryptostomata, the presence of a central vein, and the characteristics of the leaf margin, base, and apex. The shape of aerocysts and receptacles, when present, was also recorded. In addition, cross-sections of the leaves were made to determine the number of cells in the medullary region, and the type of holdfast and shape of the main growth axis were characterized. Leaves were photographed against a white background with a millimeter scale, and morphometric measurements were carried out using the software ImageJ 1.x [29]. Qualitative characteristics were evaluated through direct observation, following the morphological terminology proposed by Mattio & Payri (2011) [6]. The resulting data were used to compare morphological variation among morphotypes and to complement the results of the genetic analysis.

2.3. Amplification and Sequencing

Two organellar gene regions were amplified by PCR (Polymerase Chain Reaction): ITS-2 and COX3. For the ITS-2 region, primers 5.8S-BF [30] and 25BR-2 [31] were used, while for COX3, primers CAF4A and CAR4A [32] were employed.
PCR reactions were prepared with a final volume of 25 µL per reaction. The mix included 2.5 µL of 10× buffer, 2.5 µL of dNTPs (200 µM), 1 µL of BSA, 1 µL of forward primer, 1 µL of reverse primer, 0.1 µL of Taq polymerase, 15.9 µL of Milli-Q water, and 1 µL of DNA. Amplified products were verified by electrophoresis on 1% agarose gel and subsequently sent for bidirectional sequencing through the company Macrogen (Seoul, Republic of Korea). The resulting sequences were assembled and edited using the software Benchling (available at https://benchling.com).

2.4. Alignment and Phylogenetic Analysis

The sequences were aligned separately for the two markers using MAFFT online v.7 [33] (available at https://mafft.cbrc.jp/alignment/server/index.html, accessed on 6 March 2025), using the L-INS-i strategy, which is recommended for small alignments and sequences of varying lengths. Once aligned, the sequences were examined in MEGA 11 [34], although no manual editing was performed. Instead, the G-blocks program (v 0.91b) [35] was used to remove ambiguous regions from the alignment. The least stringent parameters were used: allowing small final blocks, permitting gap positions within the final blocks, and permitting less strict flanking positions. After removal of ambiguous regions with Gblocks, the final alignment lengths were 434 nucleotides for ITS2 and 433 nucleotides for COX3.
After editing, the online version of FindModel (available at https://www.hiv.lanl.gov/content/sequence/findmodel/findmodel.html, accessed on 6 March 2025) [36] was used to determine the best-fit nucleotide substitution model for each marker. The 28 default parameters of the program were used. One model was selected for ITS2 and three models for COX3 (one for each codon position since it is a protein-coding region). The GTR+Gamma model was identified as the best fit for both ITS2 and all three codon positions of COX3.
Phylogenetic reconstructions were carried out using Maximum Likelihood (ML) [37] in RAxML-GUI 2.0 [38]. The only parameters specified were the bootstrap substitution model (GTRGAMMA) and the exclusion of invariant sites. A total of 1000 bootstrap replicates were selected. The rest of the settings were left at default values. For COX3, a partition model was added to indicate that the region was protein-coding.
After conducting separate analyses for both markers, the results were compared to evaluate congruence. The individual ITS2 and COX3 phylogenies were generally congruent in the main clades but showed some minor differences in the resolution of certain internal nodes and relationships among closely related taxa. In both separate trees, the newly identified morphotypes formed distinct, well-supported clades, yet the level of support and the relationships with nearby species varied slightly. The concatenated tree (ITS2 + COX3) provided improved resolution and stronger support values across most nodes, clearly separating the two Costa Rican morphotypes into robust clades. Since no significant topological conflicts were found—defined as incongruence in the topology of nodes with bootstrap values (BS) greater than 80%—the two sets of sequences were concatenated. This was performed using MEGA 11 [34], resulting in a file containing both concatenated markers. This file was then subjected to a Maximum Likelihood analysis, using the parameters described above, except for the partition file, which was configured to include three partitions (the three COX3 codon positions plus the ITS2 region). Finally, the trees were visualized and edited using iTOL v6 [39].

3. Results

3.1. Morphological Analysis

A total of 67 specimens of the genus Sargassum were analyzed, collected between 2009 and 2024. Of these, 38 correspond to recent collections carried out between 2023 and 2024, 11 come from samples collected in 2009, and 18 belong to the historical collection of the Phycology Laboratory at the University of Costa Rica (UCR). Additional specimens were examined from the Luis Fournier Origgi Herbarium (USJ), the University of Michigan Herbarium (MICH), and the isotype of Sargassum liebmannii preserved at the University of California, Berkeley Herbarium (UC).
The morphological analysis revealed two clearly differentiated morphotypes, distinguished primarily by leaf characteristics, shape of the main axis, vesicle morphology, and type of holdfast (Table 1). However, the morphotypes do not fully correspond to previously reported taxonomic descriptions for the region, highlighting the need for further investigation to determine whether they represent intraspecific variation or a potentially undescribed species.
One of the morphotypes appears to fit the original description of Sargassum liebmannii (Figure 3 and Figure 4); although the isotype is incomplete, lacking both holdfast and vesicles, the remaining features are consistent with the specimens analyzed. In contrast, the second morphotype does not correspond to any previously reported descriptions for the region and exhibits a unique combination of characters that suggest its recognition as a new species (Figure 5 and Figure 6).

3.2. Phylogenetic Analysis and Congruence with Morphology

A total of 18 original sequences were obtained for the ITS-2 region and 6 for the COX3 marker (Table 2), supplemented with additional sequences retrieved from GenBank (Table A1). In total, 46 unique sequences were included in both individual and concatenated phylogenetic analyses. The ITS-2 alignment was 434 base pairs long, the COX3 alignment 433 base pairs, and the concatenated alignment reached a total of 867 base pairs, maintaining separate partitions by marker and by codon position for COX3.
Since no significant topological incongruences were found in the individual trees—defined as conflicting nodes with bootstrap support values greater than 80% the two markers were concatenated. The resulting phylogenetic tree showed a clear separation between the two morphotypes (Figure 7), which, although both grouped within the subgenus Sargassum, were placed in different sections.
The tree recovered five previously defined sections within the subgenus Sargassum: Binderiana, Polycystae, Ilicifolia, Zygocarpicae, and Sargassum. The morphotype that does not match previous descriptions was placed within the section Sargassum, suggesting that it may represent an undescribed species. The second morphotype clustered with species from Baja California, such as Sargassum sinicola and Sargassum horridum Setchell and N.L.Gardner, forming a well-supported clade that may represent a poorly characterized or yet-to-be-formally defined section within the subgenus, especially considering that no sequences of Sargassum liebmannii had been previously published.

3.3. Morphological Descriptions

  • Section Sargassum
Sargassum lacrucense Viales-Cubillo & Fernández-García, sp. nov. (Figure 3 and Figure 4).
Description: Sargassum lacrucense is characterized by bushy specimens reaching up to 44 cm in length, with well-developed discoid holdfasts. The main axis is cylindrical and smooth, with simple or repeatedly divided leaves ranging from oblong to linear in shape. Leaves have a narrow and symmetrical base and apices ranging from acute to rounded. Leaf thickness consists of 3–4 medullary cells. Margins are serrulated, and a prominent central vein is present. Petioles are short, cylindrical to flattened, and often bear spines on apical leaves. Cryptostomata are inconspicuous and arranged in rows along the central vein. Vesicles are spherical and rough, some with winged or foliated bases. Female receptacles appear in clusters along the thallus, with dichotomous branches and smooth margins, while male receptacles are solitary, branched, linear, and have smooth margins. Most leaves display cortical cell projections, which represent a distinctive feature of this morphotype. Female receptacles occur in clusters along the thallus, often arranged on dichotomously branched axes, and possess smooth margins. Male receptacles also occur along the thallus, are solitary, linear, and branched, and with smooth margins.
Habitat: This species occurs in both intertidal zones and open waters over rocky substrates. Down to 13 m depth.
Distribution in Central America: recorded along the Pacific coasts of Costa Rica, Nicaragua, and El Salvador.
Etymology: The specific epithet lacrucense honors the administrative unit canton of La Cruz, located in the province of Guanacaste, Costa Rica, where part of the type of material was collected. In addition to being one of the main localities where this species occurs, members of the local community actively participated in the collection of specimens. The community of La Cruz maintains a strong cultural and emotional connection to the sea, which adds symbolic value to the naming of the species.
Type locality: Isla Juanilla, La Cruz, Guanacaste, Costa Rica
Holotype: USJ 116864
Isotype: USJ 11685
Paratypes: USJ 22581; USJ 118666; USJ 116867; USJ 116868; USJ 116869
  • Section Sinicola/Herporhizum sect. nov. (Andrade-Sorcia) Fernández-García & Viales-Cubillo
Sargassum liebmannii J. Agardh (Figure 5 and Figure 6).
Description: Sargassum liebmannii has slender specimens that can reach up to 56 cm in length, with rhizoidal holdfasts. The main axis is terete and smooth, developing from multiple growth points arising from a prostrate basal axis that functions similarly to a rhizome. Leaves are simple, lanceolate to oblong, and often wavy. Leaf thickness consists of 8–9 medullary cells. The leaf base is rounded to narrow and typically asymmetrical, with mostly acute apices. Margins are serrated to deeply toothed, and a percurrent central vein is present. Petioles are short to medium in length and smooth. Cryptostomata are conspicuous and scattered across the leaf surface.
Vesicles are ovoid or spherical and frequently bear spines, crowns, or foliated structures. Male and female receptacles occur in clusters at the upper part of the thallus, are branched, and may have either smooth or spiny margins. A distinctive feature of this species is the presence of alternating bands of light and dark brown pigmentation on many of its leaves. Both male and female receptacles occur in clusters on branchlets that may be interspersed with leaves or entirely dedicated to reproductive structures. Receptacles are generally confined to the upper portion of the thallus and are borne on dichotomously branched axes with pointed terminal tips.
Habitat: It is found in both intertidal and open coastal waters over rocky substrates, down to 10 m depth.
Distribution in Central America: confirmed records along the Pacific coasts of Costa Rica, Panama, and Nicaragua.
Herbarium specimens: USJ 73474; USJ 73577; USJ 116870; USJ116871; USJ116872; USJ116873; USJ116874
Type locality: San Agustin, México UC-141536 (Appendix A).

4. Discussion

The results of this study confirm the presence of two morphologically and genetically distinct entities within the genus Sargassum along the Pacific coast of Central America. While one of the morphotypes corresponds to the original description of Sargassum liebmannii, the other exhibits a unique combination of morphological and molecular features that supports its recognition as a new species: Sargassum lacrucense sp. nov.
Historically, part of the Sargassum material present in Central America had been identified as Sargassum sinicola and Sargassum brandegeei. However, S. sinicola does not match either of the morphotypes described in this study, primarily due to differences in the type of holdfast, although the overall branching pattern and receptacle morphology could have caused confusion in the past [40]. Moreover, S. sinicola appears to be endemic to the Gulf of California, restricted to its northern part [11,41], a region characterized by cold-water conditions and oceanographic features such as strong tidal mixing and seasonal upwelling [42]. This pattern is not unique to Sargassum, as similar cases of endemism and restricted distribution have been documented in other algal genera such as Dictyota and Padina [43].
Similarly, Sargassum brandegeei, currently recognized as Sargassum herporhizum Setchell & N.L. Gardner [11] is also considered endemic to the Gulf of California. Although Dawson (1944) [19] and Norris (2010) [41] grouped it in the section Herporhizum along with S. liebmannii due to their shared rhizoidal holdfasts, S. brandegeei clearly differs by lacking cryptostomata [40] and is not closely related phylogenetically to Morphotype 2 (S. liebmannii) from this study, which clusters instead with other Baja California species. This provides strong evidence to exclude S. brandegeei (now S. herporhizum) from the Central American region.
Norris (2010) [41] also placed Sargassum liebmannii in the Herporhizum section, although without providing detailed information about the species. A revision of the isotype from San Agustín, Mexico, preserved at the University of Michigan Herbarium, along with the original description by J. Agardh (1847) [44], reveals that the type specimen lacks both holdfast and vesicles, which limits its interpretation. Nonetheless, the remaining morphological traits match those of Morphotype 2 described in this study. According to Andrade-Sorcia (2014) [11], S. liebmannii does not occur in the Gulf of California but is present in other parts of the tropical Mexican Pacific with conditions more similar to Central America [45]. Although those authors presented microsatellite data, no DNA barcoding sequences were available for phylogenetic comparison, so the match with Morphotype 2 was based solely on morphology, which was fully consistent.
Although Dawson and Norris [22,41] placed S. liebmannii in the Herporhizum section, the phylogenetic results of this study suggest a closer relationship with species in the Sinicola section, such as S. sinicola and S. horridum. This raises the possibility that S. liebmannii has been misclassified, and that the Sinicola section may need to be redefined using molecular evidence rather than morphological characters like holdfast type alone. The topology of our concatenated tree supports the validity of the Lapazeanum and Herporhizum sections (sensu Andrade-Sorcia, 2014 [11]) but also suggests that the current limits of the Sinicola section should be revised to potentially include S. liebmannii, or alternatively, that a new closely related section should be proposed.
The checklist of marine macroalgae from the Central American Pacific [14] also includes a record of Sargassum liebmannii var. nicoyana. However, Setchell (1937) [18] indicated that such differences could be attributed to intraspecific morphological variation. Interestingly, Setchell re-described S. liebmannii and mentioned that it had discoid holdfasts. A review of herbarium specimens from El Salvador identified as S. liebmannii by Dawson shows that they match the new species described in this study (S. lacrucense), suggesting that Setchell may have inadvertently confused the two morphotypes.
In the concatenated phylogenetic analysis, (S. lacrucense) is grouped within the Sargassum section, the same section that includes S. liebmannii, although in a separate clade. This clade also contains species such as S. spinuligerum Sonder and S. spinuligerum var. crispatum Kützing, which are distributed in the Indo-Pacific region [7,46,47] and exhibit some resemblance to the newly described morphotype, particularly in general thallus morphology [7]. However, these species differ notably in leaf and reproductive structure characteristics, suggesting that they are sister species rather than conspecifics of S. lacrucense.
Moreover, the taxonomic structure of the sections that include Gulf of California species remains partially unresolved. Although the phylogenetic tree obtained in this study supports the validity of the Lapazeanum and Herporhizum sections (in agreement with Andrade-Sorcia, 2014; Norris, 2010) [11,41], it also suggests that the Sinicola section—currently including S. sinicola and S. horridum—might also encompass S. liebmannii. This would contradict earlier proposals by Dawson and Norris, who placed S. liebmannii in Herporhizum. However, S. liebmannii does not fully match the morphological diagnosis of the Sinicola section, especially regarding its holdfast, and thus, this section may require redefinition or the proposal of a new, closely related section.
Therefore, we propose the establishment of a new section, Herporizum/Sinicola, that integrates the previously recognized sections Herporhizum, Lapazeanum, and Sinicola, based on a combination of morphological and molecular evidence. This newly proposed section is defined by the geographic distribution of its component species, which are found exclusively in the eastern Pacific, including the Gulf of California and the tropical Pacific coasts of Mexico and Central America. Morphologically, all species within this group share several key features: leaves with dentate to serrated margins, an apparent to inconspicuous central vein, and the frequent presence of cryptostomata. Despite its morphological similarities to section Herporhizum, the overall congruence between morphological traits and genetic relationships supports the unification of these three sections into a single, coherent lineage. This pattern is consistent with other genera such as Dictyota and Padina, which also include species that have diversified exclusively in the eastern Pacific [43].

5. Conclusions

This study reduces the number of Sargassum species reported in the Pacific of Central America from four to two clearly delimited entities. It also provides the first available molecular sequences for S. liebmannii and for the newly described species S. lacrucense sp. nov., representing a significant contribution to future molecular identification efforts for macroalgae in the region. We propose a new section, Herporhizum/Sinicola, uniting the former sections Herporhizum, Lapazeanum, and Sinicola based on geographical, morphological, and molecular evidence.
Based on current records, Sargassum liebmannii appears to be widely distributed across the Pacific coast of Central America, with confirmed presence in Costa Rica, Panama, and Nicaragua, although no recent records were found for El Salvador. In contrast, Sargassum lacrucense has been documented in Costa Rica, Nicaragua, and El Salvador but not in Panama. Since both species can inhabit both intertidal and subtidal rocky habitats, further sampling efforts are required to confirm their presence or absence in other parts of the region.

Author Contributions

Conceptualization, M.V.-C. and C.F.-G.; methodology, M.V.-C., C.F.-G., and K.G.-S.; software, M.V.-C.; validation, M.V.-C., C.F.-G., and K.G.-S.; formal analysis, M.V.-C.; investigation, M.V.-C., C.F.-G., F.Q.-P., P.D.-C., and K.G.-S.; resources, C.F.-G.; data curation, M.V.-C.; writing—original draft preparation, M.V.-C.; writing—review and editing, C.F.-G.; visualization, M.V.-C.; supervision, C.F.-G.; project administration, C.F.-G.; funding acquisition, C.F.-G. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Guanacaste Dry Forest Conservation Fund and by the research projects B7-903 and C3-262 registered at CIMAR (Center for Research in Marine Sciences and Limnology), Office of the Vice-rectory for Research, University of Costa Rica.

Institutional Review Board Statement

The genetic access permits were approved by the Biodiversity Committee of the University of Costa Rica.

Data Availability Statement

The original data presented in the study are openly available in GenBank repository.

Acknowledgments

We thank the administrative staff of CIMAR for their valuable support throughout this project. We are also grateful to Isabel Cordón, Kenneth Monge Barquero, Andrea Planas, Arturo Ayala, Juan José Alvarado, and Family Lara (Diving Center Cuajiniquil) for their assistance in specimen collection and sample processing and Fabián Mora Escobar for taking the photographs of the specimens in the laboratory. This research was conducted under the following research and collection permits: [R-SINAC-ACG-PI-022-2018, N° R-SINAC-ACG-PI-038-2021, N° R-SINAC-ACG-PI-019-2023, R-SINAC-ACG-PI-041-2024] in Costa Rica, [DGPN/DB/DAB-IC-0004-2009] in Nicaragua, and [AIMA-01-2009] in El Salvador. During the preparation of this manuscript, the authors used AI to assist in translating the original Spanish version of the text into English. The authors have reviewed and edited the output and take full responsibility for the content of this publication.

Conflicts of Interest

The authors declare no conflicts of interest.

Abbreviations

The following abbreviations are used in this manuscript:
MICHUniversity of Michigan Herbarium
UCThe University Herbarium of California, Berkeley
USJUniversity of Costa Rica Herbarium, Dr. Luis A. Fournier Origgi
CIBETCentro de Investigación en Biodiversidad y Ecología Tropical
CIMARCentro de Investigación en Ciencias del Mar y Limnología de la Universidad de Costa Rica
PCRPolymerase Chain Reaction
DNADeoxyribonucleic acid
ITS2Internal Transcribed Spacer 2
COX3Cytochrome Oxidase Subunit 3

Appendix A

Diversity 17 00592 g0a1
Figure A1. Isotype of Sargassum liebmannii from San Agustín, Mexico. Herbarium specimen housed at the University Herbarium, University of California, Berkeley (UC-141536) (Bar scale in cm).
Figure A1. Isotype of Sargassum liebmannii from San Agustín, Mexico. Herbarium specimen housed at the University Herbarium, University of California, Berkeley (UC-141536) (Bar scale in cm).
Diversity 17 00592 g0a1

Appendix B

Table A1. GenBank accession numbers for taxa included in the phylogenetic reconstruction of the genus Sargassum.
Table A1. GenBank accession numbers for taxa included in the phylogenetic reconstruction of the genus Sargassum.
TaxonCOX3ITS2
Cystophora retroflexaJN654446GU289346
Sargassum hystrixKP064338KM461676
Sargassum howeanumFJ170413FJ170438
Sargassum aquifoliumJN243775EU833429
Sargassum brandegeeiJX560119JX560132
Sargassum carpophyllumEU833417EU100797
Sargassum decurrensEU100822EU882257
Sargassum echinocarpumEU100835EU100795
Sargassum fallaxJN243781JN243809
Sargassum filicinumJF931740AB043108
Sargassum fluitansKP064335KM461674
Sargassum fusiformeJF931743JF931859
Sargassum herporhizumJX560118JX560130
Sargassum horneriAB430581AB043680
Sargassum horridumJX560120JX560122
Sargassum ilicifoliumEU833391EU882249
Sargassum johnstoniiJX560116JX560129
Sargassum lapazeanumJX560117JX560125
Sargassum macrocarpumJF931748KY935429
Sargassum mcclureiKY935444AB043111
Sargassum micracanthumJF931752KY935431
Sargassum miyabeiJF931753AY150007
Sargassum natansKP064340KM461678
Sargassum obtusifoliumEU100831EU100785
Sargassum pacificumEU100828EU100774
Sargassum patensJF931758AB043666
Sargassum piluliferumJF931760AB043617
Sargassum polycystumEU833405OK041991
Sargassum polyphyllumEU833402EU833424
Sargassum siliquastrumJF931765JF931843
Sargassum sinclairiiKF281971FJ170458
Sargassum sinicolaJX560121JX560124
Sargassum spinuligerumJN243785FJ170460
Sargassum spinuligerum var. crispataFJ170429FJ170463
Sargassum swartziiKF281820EU882254
Sargassum thunbergiiJF931766JF931858
Sargassum turbinarioidesEU882245EU882256
Sargassum vulgareKJ572507KJ572480
Sargassum yamadaeJF931767AB043561
Sargassum yendoiJF931771AB043667

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Figure 1. Subtidal Sargassum forests in the Eastern Tropical Pacific. (A) Dense algal cover of Sargassum liebmannii in Isla Juanilla, Costa Rica. (B) Subtidal Sargassum liebmannii forest in San Juan del Sur, Nicaragua.
Figure 1. Subtidal Sargassum forests in the Eastern Tropical Pacific. (A) Dense algal cover of Sargassum liebmannii in Isla Juanilla, Costa Rica. (B) Subtidal Sargassum liebmannii forest in San Juan del Sur, Nicaragua.
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Figure 2. Sampling sites in Central America for Sargassum analyses.
Figure 2. Sampling sites in Central America for Sargassum analyses.
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Figure 3. Morphological characteristics of Sargassum lacrucense sp. nov. (A) Habit of the species. (B) Cylindrical projections on the leaf surface. (C) Variation in leaf morphology. (D) Discoid holdfast. (E) Female reproductive structure (receptacle). (F) Morphological variation of rounded vesicles.
Figure 3. Morphological characteristics of Sargassum lacrucense sp. nov. (A) Habit of the species. (B) Cylindrical projections on the leaf surface. (C) Variation in leaf morphology. (D) Discoid holdfast. (E) Female reproductive structure (receptacle). (F) Morphological variation of rounded vesicles.
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Figure 4. Microscopic morphology of Sargassum lacrucense sp. nov. (A) Longitudinal section of the leaf at 40× magnification. (B) Longitudinal section of a cylindrical leaf projection at 10×. (C) Longitudinal section of the leaf at 10×. (D) Transverse section of the leaf at 10×.
Figure 4. Microscopic morphology of Sargassum lacrucense sp. nov. (A) Longitudinal section of the leaf at 40× magnification. (B) Longitudinal section of a cylindrical leaf projection at 10×. (C) Longitudinal section of the leaf at 10×. (D) Transverse section of the leaf at 10×.
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Figure 5. Morphological characteristics of Sargassum liebmannii. (A) Habit of the species. (B) Alternating light and dark bands on leaf blades. (C) Variation in leaf morphology. (D) Rhizoidal holdfast. (E) Reproductive structure. (F) Morphology of vesicles.
Figure 5. Morphological characteristics of Sargassum liebmannii. (A) Habit of the species. (B) Alternating light and dark bands on leaf blades. (C) Variation in leaf morphology. (D) Rhizoidal holdfast. (E) Reproductive structure. (F) Morphology of vesicles.
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Figure 6. Microscopic morphology of Sargassum liebmannii. (A) Longitudinal section of the leaf at 10× magnification. (B) Transverse section of the leaf at 10×.
Figure 6. Microscopic morphology of Sargassum liebmannii. (A) Longitudinal section of the leaf at 10× magnification. (B) Transverse section of the leaf at 10×.
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Figure 7. Maximum Likelihood phylogenetic tree based on concatenated ITS2 and COX3 sequences of the genus Sargassum. Bootstrap support values (ML) are shown at the center of the nodes (in boxes); only values above 50% are displayed.
Figure 7. Maximum Likelihood phylogenetic tree based on concatenated ITS2 and COX3 sequences of the genus Sargassum. Bootstrap support values (ML) are shown at the center of the nodes (in boxes); only values above 50% are displayed.
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Table 1. Taxonomic comparison of Sargassum lacrucense (sp. nov.) and Sargassum liebmannii based on vegetative and reproductive characters.
Table 1. Taxonomic comparison of Sargassum lacrucense (sp. nov.) and Sargassum liebmannii based on vegetative and reproductive characters.
CharacteristicS. lacrucense (sp. nov.)S. liebmannii J. Agardh
Specimen sizeUp to 44 cm, shrubbyUp to 56 cm, slender
Type of holdfastDiscoidRhizoidal
Main axis shapeCylindrical and smoothTerete, smooth, grows from rhizoid
Leaf shapeSimple or divided multiple times; oblong to linearSimple, lanceolate to oblong, mostly undulate
Leaf dimensions (cm) (L/W)1.21–6.23/0.54–21.27–9.05/0.37–1.40
Leaf marginsSerrulateSerrulate to deeply dentate
Central veinPercurrentPercurrent
PetioleShort, cylindrical to flattened, with apical spinesShort to medium, smooth
Leaf baseNarrow and symmetricalRounded to narrow, asymmetrical
Leaf apexFrom acute to roundedMostly acute
Leaf thickness3–4 medullary cells8–9 medullary cells
CryptostomataInconspicuous, in rows along the veinConspicuous, scattered
VesiclesSpherical, rough, some with foliar-like basesOvoid or spherical, majority with spines
Receptacles (♀)In clusters along the thallus, with dichotomous branches and smooth marginsIn clusters on upper thallus, with smooth or spiny margins
Receptacles (♂)Solitary, branched, linear with smooth marginsIn clusters on upper thallus, branched with smooth margin
RemarksLeaves with conical projections on surfaceLeaves with alternating light/dark brown bands
Systematic positionSubg. Sargassum, sect. SargassumSubg. Sargassum, sect. Sinicola/Herporhizum
Table 2. Origin and GenBank accession numbers of new Sargassum sequences obtained for this study.
Table 2. Origin and GenBank accession numbers of new Sargassum sequences obtained for this study.
TaxonVoucherCollection SiteITS2COX3
Sargassum liebmanniiUCR-2161Playa Piedras, El Jobo, La Cruz, Guanacaste, Costa RicaPX060136 PX060137-
Sargassum liebmanniiUCR-2172Isla Muñecos, Bahía Salinas, La Cruz, Guanacaste, Costa RicaPX060138-
Sargassum liebmanniiPAN-244Las Perlas, Golfo de Panamá, Panamá-PX214380
Sargassum liebmanniiNIC-349La Flor, San Juan del Sur, Nicaragua-PX214378
Sargassum liebmanniiUCR-2201Bajo Rojo, Cuajiniquil, La Cruz, Guanacaste, Costa Rica-PX214379
Sargassum lacrucenseUCR-2196Isla Muñecos, Bahía Salinas, La Cruz, Guanacaste, Costa RicaPX060141-
Sargassum lacrucenseUCR-2174Playa Piedras, El Jobo, La Cruz, Guanacaste Costa RicaPX060139PX214380 PX214381
Sargassum lacrucenseNIC-348La Flor, San Juan del Sur, Nicaragua-PX214382
Sargassum lacrucenseUCR-2131Isla Pelada, Islas Murciélago, Guanacaste Costa RicaPX060140-
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Viales-Cubillo, M.; Quesada-Perez, F.; Díaz-Canales, P.; González-Sánchez, K.; Fernández-García, C. Integrative Approach to Species Delimitation in Sargassum (Fucales, Phaeophyceae) from Central American Pacific Based on Morphological and Genetic Evidence. Diversity 2025, 17, 592. https://doi.org/10.3390/d17090592

AMA Style

Viales-Cubillo M, Quesada-Perez F, Díaz-Canales P, González-Sánchez K, Fernández-García C. Integrative Approach to Species Delimitation in Sargassum (Fucales, Phaeophyceae) from Central American Pacific Based on Morphological and Genetic Evidence. Diversity. 2025; 17(9):592. https://doi.org/10.3390/d17090592

Chicago/Turabian Style

Viales-Cubillo, Mariana, Fabio Quesada-Perez, Paola Díaz-Canales, Kaylen González-Sánchez, and Cindy Fernández-García. 2025. "Integrative Approach to Species Delimitation in Sargassum (Fucales, Phaeophyceae) from Central American Pacific Based on Morphological and Genetic Evidence" Diversity 17, no. 9: 592. https://doi.org/10.3390/d17090592

APA Style

Viales-Cubillo, M., Quesada-Perez, F., Díaz-Canales, P., González-Sánchez, K., & Fernández-García, C. (2025). Integrative Approach to Species Delimitation in Sargassum (Fucales, Phaeophyceae) from Central American Pacific Based on Morphological and Genetic Evidence. Diversity, 17(9), 592. https://doi.org/10.3390/d17090592

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