Research Article |
Corresponding author: Carlos Sánchez ( csanchez@uabcs.mx ) Academic editor: James Reimer
© 2023 Osvaldo Hernández, Jaime Gómez-Gutiérrez, Carolina Galván-Tirado, Carlos Sánchez.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hernández O, Gómez-Gutiérrez J, Galván-Tirado C, Sánchez C (2023) Three new species of the sea fan genus Muricea (Cnidaria, Octocorallia, Plexauridae) from the northwest region of Mexico. ZooKeys 1169: 333-352. https://doi.org/10.3897/zookeys.1169.89651
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Twenty-one nominal species of Muricea have been reported in the Eastern Pacific with nine of them reported in the Mexican Pacific. We describe three new species of Muricea: Muricea ambarae sp. nov. and Muricea cacao sp. nov., from rocky reefs on the central and the northern Gulf of California and the Pacific coast of Baja California Sur, and Muricea molinai sp. nov., from the Pacific coast of Baja California Sur. Muricea ambarae sp. nov. and M. cacao sp. nov. are taxonomically allied to the nominal species Muricea fruticosa Verrill, 1869 due to the morphological similarity of colony growth patterns and the phylogenetic closeness based on the mitochondrial MutS gene (mtMutS); but differ mainly in the calyx form and composition of sclerites. The main morphological differences between the new Muricea species are in their sclerite forms and color; M. ambarae sp. nov. has orange-colored colonies, thin leaf spindles and tuberculated blunt spindles, while M. cacao sp. nov. has dark brown colored colonies, strong spinous spindles and an absence of tuberculated blunt spindles. Muricea molinai sp. nov. is phylogenetically close and morphologically similar to Muricea squarrosa Verrill, 1869 in the growth form of the colony and tubular calyces; but has dark brown colored colonies and has calyces from the base to the branch tips. With these three new species, the total number of Muricea species reported in the Mexican northwest region increases to twelve and a total of 24 nominal species in the Eastern Pacific.
Baja California, Gulf of California, Malacalcyonacea, mtMutS, species-groups, taxonomy
Sea fans of the family Plexauridae of the genus Muricea Lamouroux, 1821 currently include 30 nominal species worldwide: nine species are distributed in the Western Atlantic Ocean and 21 species are distributed in the Eastern Pacific (
Nine Muricea species have been reported on the Gulf of California and Pacific coast of Mexico: Muricea austera Verrill, 1869; Muricea californica Aurivillius, 1931; Muricea echinata Verrill, 1866; Muricea formosa Verrill, 1869; Muricea fruticosa Verrill, 1869; Muricea hebes Verrill, 1864; Muricea plantaginea (Valenciennes, 1846); Muricea purpurea Verrill, 1868; and Muricea robusta Verrill, 1864 (
All sea fan colonies analyzed for the present study were collected by scuba diving (< 50 m depth) during systematic monitoring surveys carried out between 1998 and 2020 in 250 locations in the Gulf of California and along the Pacific coast of Baja California Peninsula, Mexico (Fig.
Locations of sample collections of the three new species of the genus Muricea discovered along the Pacific coast of Baja California Sur and the Gulf of California, Mexico A scuba diving and monitoring census sites B Muricea ambarae sp. nov. collection sites C Muricea cacao sp. nov. collection sites D Muricea molinai sp. nov. collection sites.
Sea fan colonies preserved dry were used for molecular analyses and ten sequences (six sequences from the three new species, plus four sequences of nominal species) of a partial fragment (~931 base pair, bp) of the mtMutS gene were generated. The sequences were submitted to GenBank with accession numbers given in Suppl. material
Phylum Cnidaria Hatschek, 1888
Class Anthozoa Ehrenberg, 1834
Subclass Octocorallia Haeckel, 1866
Order Malacalcyonacea McFadden, van Ofwegen & Quattrini, 2022
Family Plexauridae Gray, 1859
Holotype. USNM 1606629: dry, San Esteban Island (Punta Sureste), Sonora, Mexico (28°40.29228'N, 112°33.24035'W), 20 m depth, 20 °C, 20 June 2010 (Fig.
Colony with flabellate growth in one plane and laterally branched, 21 cm high and 14.2 cm wide (Fig.
Internal and external characters of Muricea ambarae sp. nov., Muricea cacao sp. nov. and Muricea molinai sp. nov. with similar Muricea nominal species distributed along the Mexican Pacific and Gulf of California collected from 1998–2020 and compared with
Species | Colony growth | Branching type | Terminal branches length (cm) | Polyp distribution rows | Pseudoanastomosis | Calyx height elevation (mm) | Calyx form | Colony Color | Outer coenenchymal and calyx dominant spindles | Coenenchymal and calycular spindles maximum size (mm) | Inner coenenchymal spindles | Anthocodial sclerites | Sclerites color |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
M. californica | bu | irr, lb | 2.8 | c, im | no | 1.9 | el | ro | ls | 0.5 | slr, ws | lb, wr | am, lo, ro, py |
M. echinata | bu | irr, lb | 6–3 | c | no | 2.8–3 | sl | rb | uss | 2.4 | ws | r, bs | o, lb |
M. fruticosa | bu | irr | 1.5–4 | c | no | 1–1.2 | sl | rb, w, bi | uss | 2 | ws | ws, wr, bs | w, rb, py |
M. galapagensis | fa | ob | 8 | s | no | 0.6–1 | sl | lo | uss | 4.1 | ws | r, ps | am, lo |
M. plantaginea | fl | irr, lb | 1–5 | c, im | no | 0.7–1.2 | sl | db/w | ls | 1 | ws | lb, wr | rb, am |
M. squarrosa | fl | di | 4 | c | no | 2.6 | t | lb | cl, cs | 1.3 | tbs | cl | br, cl, py, y |
M. ambarae sp. nov. | fl | lb | 9.5 | im | no | 1 | sl | o | ls, tbs | 1.2 | ws | lb | cl, o, y, py |
M. cacao sp. nov. | fl | lb | 9 | im | yes | 1 | sl | br-r | ls | 1.7 | ae/de-ws | tbr-ae | db, rb |
M. molinai sp. nov. | fl | lb | 8.7 | im | no | 3 | t | g | uss, tbs | 2.5 | str, cs | tbr, lr | g, am, cl |
The sclerites of the outer coenenchyme and calycular are pale yellow or pale orange leaf spindles (0.3–1.2 mm length), tuberculated spindles with blunt ends (0.2–1.1 mm length), and tuberculated spindles with acute ends (0.2–1.1 mm length) (Figs
All 14 colonies of Muricea ambarae sp. nov. examined are morphologically like the holotype in colony growth and sclerite form with colony size range observed in situ between 4 and 32 cm height (Suppl. material
Muricea ambarae sp. nov. was collected at two locations, the northern and central regions of the Gulf of California (where the species is more frequently collected), and Bahía Magdalena and Punta Abreojos located along the Pacific coast of Baja California Sur, Mexico (Fig.
Muricea ambarae sp. nov. (Fig.
The word “ambarae” means “amber”, a hard, transparent, fossilized resin produced by some trees. Amber has colorations from pale yellow/orange to a dark orange, like the coloration observed in living colonies of Muricea ambarae sp. nov. Mexican amber, also known as Chiapas Amber, dates from 15 to 23 million years old. Since the time of the Mayan culture, its people have believed amber to have healing and protective qualities. The species name is also inspired from the name of the daughter (Ámbar) of Carlos Sánchez.
Holotype. USNM 1606633: dry, San Marcos Island (El Faro-Lobera), Baja California Sur, Mexico (27°15.95706'N, 112°5.51208'W), 15 m depth, 23 °C, 14 July 2010 (Fig.
The holotype is a brownish-red colony, growing flabellate upwards in one plane and laterally branched, 15.8 cm tall and 8.7 cm wide (Fig.
The sclerites of the outer coenenchyme and calyces are dark brown or reddish-brown (Fig.
All the Muricea cacao sp. nov. specimens collected and observed in situ are morphologically consistent with the macro- and micro-morphology of both the holotype (Fig.
Muricea cacao sp. nov. is present in the northern part of the Gulf of California from the Santa Rosalía region to the Midriff Archipelago Region (MAR) and along the Pacific coast of Baja California Sur, between Todos Santos and Bahía Magdalena (Fig.
Paratypes of Muricea cacao sp. nov. have low morphological variability and the main difference is the width of the branches (Suppl. material
The latinized species name “cacao” comes from the ancient pre-hispanic Nahuatl “cacao”, which is the seed used in the making of chocolate, but in Latin is “cacao”, the species name of the cocoa tree is Theobroma cacao L. This “chocolate” color is a practical diagnostic characteristic of both preserved colonies and live colonies with retracted polyps, distinguish this species in situ from other Muricea species.
Holotype. USNM 1606637: dry, Todos Santos, Punta Lobos (Bajo Fondo del Medio), Pacific coast of Baja California Sur, Mexico (23°21.34806'N, 110°15.40374'W), 35 m depth, 19 °C, 27 August 2016 (Fig.
Muricea molinai sp. nov. colony coloration is gray with dark gray calyces and a creamy light gray coenenchyme (Fig.
The sclerites of the outer coenenchyme and calyx are unilateral (weakly) spinous spindles and tuberculated spindles (1.2–2.5 mm in length) (Figs
All twelve Muricea molinai sp. nov. colony specimens collected, including the paratypes ranging between 7 and 21 cm height (Fig.
Muricea molinai sp. nov. is absent in the Gulf of California but overlaps its distribution and habitat with Muricea ambarae sp. nov. and Muricea cacao sp. nov. along the southwest coast of the Baja California Peninsula (Fig.
Live colonies of M. molinai sp. nov. (Fig.
Muricea molinai sp. nov. is named in memory of Dr. José Mario Molina Pasquel y Henríquez (1943–2020), the first Mexican in 1995 to win the Nobel Prize in Chemistry. Molina played a vital role in the discovery of the Antarctic ozone hole demonstrating that chlorofluorocarbon gases were the cause of the deterioration of the ozone layer.
For several sea fan genera, the mtMutS gene is conserved to species level; but it is divergent enough to discriminate among genera and species-groups in octocorals. The partial fragment used for the phylogenetic reconstruction was highly conserved in Muricea, nonetheless and even with a lack of strong node support, Muricea ambarae sp. nov. and Muricea cacao sp. nov. were separated in a well-defined clade together with the nominal species Muricea fruticosa. In the case of Muricea molinai sp. nov., although not falling into a clade and showing unresolved relationships, it is clearly distinct from these last two new species and close to the nominal species Muricea squarrosa and Muricea hebes. The comparison of these mtMutS sequences with M. plantaginea, M. californica and M. squarrosa support the morphological evidence that, M. ambarae sp. nov., M. cacao sp. nov. and M. molinai sp. nov. are new species (Fig.
Sea fan species assemblages have a clear latitudinal regionalization in the Gulf of California (
Maximum-likelihood phylogenetic reconstruction of partial mtMutS gene for Muricea reported for the Pacific Ocean showing in bold the sequences of the present study of Muricea ambarae sp. nov., Muricea cacao sp. nov. and Muricea molinai sp. nov. Numbers above the branches are bootstrap percentage values.
The molecular analysis of species of the genus Muricea and other sea fan genera distributed in the Eastern Pacific is still incipient (
We thank the scientists, owner and crew of the liveaboard “Rocío del Mar” and “Quino El Guardián” for their valuable help during the scuba diving collections. We especially thank Ariel Cruz-Villacorta, Scanning Electron Microscopy laboratory manager at the Centro de Investigaciones Biológicas del Noroeste S.C. (CIBNOR). Thanks to Juan Carlos Hernández Meijueiro for his valuable help in defining the correct etymology for the naming of the new species. We are grateful for the research collection facilities and permission of Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT) and Comisión Nacional de Áreas Naturales Protegidas (CONANP), Mexico (F00-0526). Universidad Autónoma de Baja California Sur and Centro Interdisciplinario de Ciencias Marinas, Instituto Politécnico Nacional through the Project Taxonomy of Octocorals of the genus Muricea in the Gulf of California (SIP-IPN 20220094) provided additional financial support. This research was conducted as part of the Universidad Autónoma de Baja California Sur, Proyecto Fauna Arrecifal (
No conflict of interest was declared.
No ethical statement was reported.
No funding was reported.
Conceptualization: CS, OH, JGG. Investigation: CGT, CS, JGG, OH.
Osvaldo Hernández https://orcid.org/0000-0001-6195-149X
Jaime Gómez-Gutiérrez https://orcid.org/0000-0003-2516-897X
Carolina Galván-Tirado https://orcid.org/0000-0003-3973-256X
Carlos Sánchez https://orcid.org/0000-0001-7199-1268
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary data
Data type: morphological, images, molecular data
Explanation note: We showed the paratypes images of the three sea fans described species. Also, the in situ images comparison between the described species with other nominal species of Muricea. The methodology of the extraction and molecular analysis is included, and the table of accession numbers in GenBank.