©The
World Botanical Associates Web Page
Prepared by Richard W. Spjut
April 2003, Oct. 2005, Sep 2012, Sep 2021,
Dec 2021, Aug 2022
Niebla and Vermilacinia (Ramalinaceae) from California and Baja
California.
Evolutionary history of coastal species
of fog lichen genera
Spjut R, Simon A, Guissard M, Magain N, Sérusiaux E.
2020. The fruticose genera in the Ramalinaceae (Ascomycota, Lecanoromycetes):
their diversity and evolutionary history. MycoKeys. 73: 1–68.
published online.
Evolution and diversification of Niebla
Additional Discussion and References: See: Introduction to Niebla and its phylogeography
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Bahía de San Quintín, |
Monterey County, |
Marin County, |
Bahía de San Quintín,
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TLC profile for selected species of Niebla received from Dr. Harinantenaina Liva Romuald Rakotondraibe at the Ohio State University, Oct 2017. Niebla eburnea L2 (17805) and L10 (17811). N. eburnea from the Monterey Peninsula (L10) shows two sky blue spots for Godin Reagent absent in the N. homalea and the other N. eburnea specimen from Stinson Beach, Marin Co. This may be due to differences in triterpenoids. Divaricatic acid shown for both specimens. N. disrupta was reidentified N. homalea based on TLC. |
Niebla eburnea ITS and six loci phylogenetic trees in part showing two separate clades with N. homalea in California and in Baja California. Baja specimens collected near Bahía Falsa west of San Quintín. California specimens shown above associated with N. homalea, 17811 from the Monterey Peninsula most closely related to 17805 on the Pt. Reyes Peninsula south of Stinson Beach. |
Bahía de San Quintín, |
San José Ranch between Punta Canoas and Punta Blanca,
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Puerto Catarina, Spjut |
Puerto Catarina, Spjut |
Cerro Solo, BCN, Spjut & |
San Andrés Ranch, |
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Above Point Pedernales, CA |
Niebla eburnea is a common fruticose lichen on rocks along the California coast from Mendocino Co. to San Diego Co., occurring also in the Channel Islands, and less frequently further south in Baja California to near Punta Santa Rosalillita. The original (type) collection was from a rock outcrop within ocean mist near Cerro Solo along the Pacific Coast of Baja California between San Vicente and Colonet. It was part of sample submitted for anti-HIV screening that was collected on the vertical face of a isolated 1.5 tall rock on a beach in association with other fog lichens, N. josecuervoi, Vermilacinia laevigata, V. ceruchoides, and V. procera. Duplicates were later provided to the Lichen Exchange at ASU, in addition to isotypes that may have been distributed by the Smithsonian Institution (intended for BCMEX, LA). The type collection was reported to have divaricatic acid but later found to actually have isodivaricatic acid as determined by a natural product chemist at Ohio State University (Professor Rakotondraibe) employing NMR. Niebla eburnea is identified by the ± creamy (ivory)-glazed cortex (skin-like or like frosting on a pastry), by its branches half-twisted near apex and near base—in which the margins ± do not alternate (cross or unite), by the terminal part of a main branch often digitately divided into finger-like or claw-like branchlets (fringing branchlets), and by the key lichen substance, divaricatic acid (with triterpenes and basal pigment skyrin), or isodivaricatic acid from the type collection collected just west of Cerro Solo, Baja California in May 1985. The type for Niebla eburnea was collected near Colonet, BC. It resembles the type for N. cornea (sekikaic acid) from Morro Bay in having mostly simple, apical dilated branches uniformly blackened around base (from pigment skyrin). The two species are distinguished by their chemistry. Vermilacinia laevigata can be easily confused with these species. It differs by the absence of chondroid strands, and by the terpenes zeorin (triterpene) and [-]-16 α-hydroxykaurane (diterpene); N. eburnea has triterpenes nieblastictanes and nieblaflavicanes. Occasionally N. eburnea is difficult to distinguish from N. homalea (divaricatic acid) and N. disrupta (sekikaic acid), which generally have narrower branches that twist 90° at frequent intervals between the base and apex, and which have a uniformly glossy cortex transversely cracked at various intervals. Problem thalli have ±uniformly narrow branches with a transversely cracked cortex (e.g., Bratt 5591 from Point Pedernales, CA, photo 10.3, Spjut 1996). DNA phylogeny shows these in sister clades with N. homalea cryptic species, and also N. testudinarai; thus, phylogenetically unresolved based on phenotypic identifications. The phylogeny for N. eburnea in Spjut et al. (2020) included four specimens from three locations, two in northern California (Monterey Peninsula , Pt. Reyes) and one in Baja California near San Quintín for two specimens. Analyses by BPP and Stacey both delimited three species that likely correspond to three clades. In the phylogenetic tree (Fig. 7), the species in Baja California is monophyletic. In California, the species is not separated from N. homalea in which both occur together in two separate clades. Thus, in northern California both N. homalea and N. eburnea comprise cryptic species, while N. eburnea may also be viewed as a cryptic hybrid (or allasomorph), phenotypically different but genetically undifferentiated from N. homalea. As noted above, the type collection reportedly contains only isodivaricatic acid (Professor Rakotondraibe, pers. comm.), in contrast to divaricatic acid confirmed for specimens collected of N. homalea at the putative type locality at Pt. Reyes (Zhang et al. 2020). Whether cryptic species relate to differences in isomers remains to be determined while it may be further noted that they can have different physical properties; for example, isodivaricatic acid has shown activity against a cancer cell line, but not divaricatic acid, although reported in literature to have antifungal and antibacterial activities. The related N. testudinaria differs by the regular appearance of reticulate cortical ridges resulting in the branches appearing more prismatic than sub-tubular; it occurs from California to Baja California Norte, especially in the Channel Islands, and is also on Isla Guadalupe and on Isla San Martín. Niebla testudinaria was found in a separate clade from N. eburnea, but not from N. homalea (Spjut et al. Fig. 7). Their differences are summarized in the following table, the bold type indicating the diagnostic features.
Niebla eburnea includes variation that might be recognized as distinct varieties but here considered 'variants' as outlined below with reference to geographic locations from where specimens were collected.
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