Niebla fimbriata

©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
April 2003, Oct. 2005, Sep 2012
Additions May 2017, Dec 2021

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
, Ramalina and Vermilacinia
Emmanuel Sérusiaux & Richard  Spjut
Baja California, Jan-Feb 2016

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. MycoKeys. 2020;73:1-68. Published 2020 Sep 11. doi:10.3897/mycokeys.73.47287

Evolution and diversification of Niebla
Steve Leavitt et al., Baja California, Dec 2016

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.

Introduction to Niebla



Niebla fimbriata typical

San Antonio del Mar, 92 m, Spjut & Sérusiaux 17012-4670. Jan 2016. Note differences in triterpenes for three sekikaic acid specimens, which include trace of divaricatic acid.. Image for 17009-4669 shown below, morphologically atypical. 17021-4676 is N. palmeri.

Southwest of San Quintín in the Punta Mazo Reserve, on volcanic slopes of Volcan Sudoeste, 40 m, Leavitt et al. 16-716, Dec 2016

Laguna and peninsula of San Quintín, N 30°29'743, W 116°00.044', 4 m, Spjut & Sérusiaux 17032B-4677,. Jan 2016

Just north of the city of El Rosario, along the Transpeninsular Hwy 1, on west side of road on small ridgeline dominated by euphorbs and small rocks, 150 m, Leavitt et al. 16-821, Dec 2016

Niebla aff. fimbriata

San Antonio del Mar, 92 m, Spjut & Sérusiaux 17009-4669. Jan 2016. .


West of San Telmo, along road, Leavitt et al. 16-1098, Dec 2016



fimbriata-11154.jpg (132384 bytes)

Punta Blanca, Spjut & Marin 11454, Apr 1990


fimbriata-11177.jpg (48563 bytes)

San Antonio del Mar, Spjut & Marin 11177, Apr 1990

San Antonio del Mar, Spjut & Marin 10503, Apr 1989

fimbriata-13031.jpg (106821 bytes)

Punta Canoas, Spjut & Marin 13031, Apr 1994

Punta Canoas, Spjut & Marin 13079, Apr 1994

Punta Canoas, Spjut & Marin 13111, Apr 1994

San Antonio del Mar,  Apr 1990

San Antonio del Mar, Spjut 10232, Mar 1988


SW of el Rosario along
road to Punta Baja,
Spjut 10288b, Mar 1988

Cañón San Fernando, BCN, Spjut & Marin 12729, March 1993

Santa Cruz Island, Bratt 6436

Arroyo Sauces just S of Punta Cuchillo between Punta Canoas and Punta Blanca, Spjut & Marin 11448, April 1990

Geographical Distribution

Portion of a Niebla phylogenetic ITS tree, including sequences for Spjut et al. (2020) reviewed for Jorna et al.. (2021) within the large sekikaic acid clade; identification changes and geographical data added by R. Spjut.

San Antonio del Mar,  Apr 1990, Spjut 11184, holotype

Portion of ITS phylogenetic tree showing N. fimbriata and N. palmeri species complexes generated by Professor Sérusiaux for Spjut et al. (2020)

     Niebla fimbriata is a fruticose lichen occurring infrequently along the Pacific  Coast of the Baja California peninsula from Punta Blanca north to San Antonio del Mar and the Channel Islands in California. It appears most common in the Chaparral-Desert-Transition (CDT) between San Quintín and Cabo Colonet, especially on lava rubble of mesas such as east of San Antonio del Mar, the type locality where it may be found with depsidone species, N. josecuervoi (salazinic acid), N. effusa (salazinic acid), and N. pulchribarbara (protocetraric acid).  

     Niebla fimbriata is characterized by having sekikaic acid with triterpenes, occasionally with both sekikaic acid and  divaricatic acid, and by the wind-swept spine-like (fimbriate or fringed) branchlets along a primary branch, all spreading nearly at right angles, ± in the same direction.  The fragmentation branchlets are rather brittle, easily breaking off near the main branch. The fragile cortex is often dark olive green with frequent dimples or ripples, in contrast to the leathery, turgid, pale yellowish green cortex of N. suffnessii without basal dark pigmentation and without differentiation of fragmentation branchlets. 

     Specimens of Niebla fimbriata from the Channel Islands were included by their pinnatifid branching but differ by less defined  branch margins with the intra-marginal cortical surface appearing more strongly rugose. These features appear to intergrade with Channel Island thalli identified  N. siphonoloba, a species generally distinguished by its small tufts of simple tubular basal branches.  Judging from DNA phylogeny of the Baja California specimens, the Channel Island specimens are likely to belong to other species, possibly N. dactylifera, currently regarded endemic to San Nicolas Island.  Thalli collected near Punta Canoas appear to intergrade with N. suffnessii.

     In Baja California, Niebla fimbriata often appears similar to N. juncosa var. spinulifera, which differs by having divaricatic acid. Its branchlets appear less differentiated, ascend rather than diverge widely from the main branch.

     The phylogeny of Niebla fimbriata is problematic by appearing polyphyletic in relationship to other phenotypic species  included (associated) within its sekikaic acid group and sister groups. Species repeatedly associated with N. fimbriata are N. lobulata, N. aff. palmeri, N. palmeri, and N. siphonoloba In view of Spjut et al. (2020) and Jorna et al. (2021), they can be  referred to as species complexes by their phylogeographic relationships. For example, in the CDT, they have been collected together near Punta Colonet, San Telmo, San Antonio del Mar, San Quintín, and Punta Baja (Spjut et al. 2020; Jorna et al. 2021).  Both BPP and Stacey methods applied by Spjut et al. 2020 (Suppl. files S 3, S5) delimited two species for N. fimbriata and two for N. palmeri, four for N. lobulata, and one each for N. siphonoloba and N. suffnessii; the latter two represented by specimens only from their  type locality near Cerro Elephante on the Vizcaíno Peninsula.  In view of their phylogeny in Spjut et al. (2020, Fig. 7),  combined with data in Jorna et al. (2021), which included seven topotypes for N. lobulata, in three clades, and one additional topotype in a sister clade of unresolved species and another sister subclade with one specimen from the Vizcaíno Peninsula  sister to N. suffnessii, these  are cryptic species complexes within which cryptic species can be recognized in those phylogenies that include topotypes. 

     In a narrowest sense,  N. fimbriata appears supported by a topotype, 17012-4670 (Spjut et al. Fig. 7), if epitypified by that specimen from San Antonio del Mar since it appears  monophyletic in a basal subclade sister to subclade of sekikaic acid species complex (N. aff. fimbriata, atypical N. lobulata (not from type locality), N. palmeri)   within a "terminal" sekikaic acid clade to a divaricatic acid clade.  All specimens differ in their triterpenes. However, BPP and Stacey analyses may  not support this division, only a broader subdivision (larger clades) in which N. aff palmeri replaces N. palmeri in the association of cryptic species. Niebla aff. fimbriata 17009 also appears to be a hybrid.

     It remains to be determined whether atypical specimens identified N. siphonoloba and N. palmeri can be distinguished from N. fimbriata by DNA to correspond to their phenotypic characteristics.  For now they appear as cryptic hybrids, or allasomorphs, species that exhibit different morphotypes.  It may be noted that N. flagelliforma (divaricatic acid species) in Fig. 7 of Spjut et al. (2020) is identified N. fimbriata in their Suppl. data S3 where correctly identified. 

For more discussion and reference materials see Introduction to Niebla