Niebla and Vermilacinia (Ramalinaceae) from California and Baja California.  
Spjut, R.W., 1996. ISSN 0833-1475, 208 pp.  
Sida, Botanical Miscellany: 14. Botanical Research Institute of Texas, Inc.
 

Evolutionary history of coastal species of fog lichen genera
Niebla, Ramalina and Vermilacinia
Emmanuel Sérusiaux & Richard  Spjut
Baja California, Jan-Feb 2016
 

Spjut R, Simon A, Guissard M, Magain N, Sérusiaux E. The fruticose genera in the Ramalinaceae (Ascomycota, Lecanoromycetes): their diversity and evolutionary history.  MycoKeys. 2020 Oct 30;74:109-110].
MycoKeys. 2020;73:1-68. Published 2020 Sep 11. doi:10.3897/mycokeys.73.47287

Jorna J, J Linde, P Searle, A Jackson, M-E Nielsen, M Nate, N Saxton, F Grewe, M de los Angeles Herrera-Campos, R Spjut, H Wu, B Ho, S Leavitt, T Lumbsch.  Species boundaries in the messy middle -- testing the hypothesis of micro-endemism in a recently diverged lineage of coastal fog desert lichen fungi. Ecology and Evolution. Published Online: 20 Dec 2021.  https://onlinelibrary.wiley.com/doi/full/10.1002/ece3.8467.

Additional Discussion: See: Introduction to Niebla and its phylogeography

NW of  Bahía de Asuncion, near the Mesa El Elephante, N 27°20.732, W 114°25.384, 120-125 m, Spjut & Sérusiaux 17145 and 17149. Jan 2016

ITS for N. rugosa, in groups
16, 17 and 18.

Isla Cedros,
Spjut & Marin 10539,
Apr 1989

     Niebla rugosa is a species of fruticose lichen endemic to the Pacific Coast Region of Baja California, infrequently occurring from the Vizcaíno Peninsula north to near Punta San Carolos, and also on Isla Cedros.  It is found on boulders on Mesa Camacho, rock walls in a narrow wind-sheltered canyon on the eastern side of Isla Cedros, on pebbles near San Carlos, and on gypsum near Bahía Tortugas; the locations are generally further away from the ocean than other related species.  The species is distinctive for the ladder-like cortical ridges on tubular branches.

     Niebla rugosa resembles N. podetiaforma in the small tufts of subtubular branches, but differs in the branches sharply angled along margins, in contrast to being rounded in N. podetiaforma. Transverse cortical ridges that connect between the margins are ± parallel  like a step ladder. As the epithet implies, Niebla rugosa has a wrinkled thallus due in part to the prominent transverse cortical ridges.  Niebla podetiaforma cortical ridges divide between margins.  Additionally, branches of N. rugosa are closely undulate along margins, especially upper third to upper half in which the marginal indentations form lobules (phyllidia). Phyllidia are absent in N. podetiaforma.  Phyllidia may be replaced by apothecia as seen in Leavitt et al. 16-915, 16-1006.

     Another species with small tufts of mostly simple basal branches, Niebla siphonoloba, differs in the cortical ridges interconnecting between margins, which seems related to the prismatic outline of the branches, although some areas of a branch may have undivided transverse connecting ridges. Generally, N. rugosa has wide rectangular areas between cortical ridges, in contrast to circular to square areoles in N. siphonoloba.   They also differ in chemistry, divaricatic acid (with triterpenes) in N. rugosa; sekikaic acid (with triterpenes) in N. siphonoloba. 

     Another distinctive feature of N. rugosa is the cupular apothecia on saucer-like marginal lobes.  The apothecia are perpendicular to the saucer lobe in contrast to slightly elevated  otherwise similar apothecia seen in N. contorta.  

    Niebla rugosa is supported by ITS phylogeny Group 18 (6-loci in Spjut et al. 2020, 17145-4768) from the Vizcaíno Peninsula north of Cerro Elephante on a mesa bordering Arroyo  San Andrés based on four specimens (Spjut & Sérusiaux 17145, 17146, 17149, 17151)  as shown above.  The type locality is northwest of Bahía Tortugas. They occur in a subclade within a large divaricatic acid group of species characterized by inflated to slightly inflated  tubular basal branches (N. contorta,  N. podetiaforma, N. turgida, N. undulata) and N. flagelliforma.  The 6-loci phylogeny shows N. rugosa in a basal position where most closely related to specimens collected in Chaparral Desert Transition rather than to those specimens collected from the Northern Vizcaíno Desert (NVD). A combined phylogeny for Jorna et al. (2021) shows one specimen, 16-1006,  collected in the southern NVD (Morro Santo Domingo) unresolved in a terminal subclade with the same species mentioned above but somewhat of a reverse relationship with basal clades represented by specimens collected in Coastal Sage Chaparral and Chaparral Desert Transition regions.

Additional References: See Niebla.