Niebla arenaria

©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
April 2003.  Comments and illustrations, Oct. 2005, Sep 2012
Additions: May 2017, Dec 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
Manuscript presented 2021 on Authorea. April 05, 2021.

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. .

Additional Discussion: See: Introduction to Niebla and its phylogeography


Punta Morro Santo Domingo, ~40 km N of Guererro Negro, solitary thallus loose on sand. Spjut & Sérusiaux 17254
Jan 2016

Punta Canoas, Leavitt et al. 16-969
Dec 2016

Bahía de San Quintín,
Laguna and peninsula

Spjut & Sérusiaux 17052
Jan 2016


Bahía de San Quintín,
Laguna and peninsula
Spjut & Sérusiaux 17050-4692
Jan 2016


South of El Rosario along road to Punta Baja, on sandy, wind-swept ridgeline. Leavitt et al. 16-1005


arenaria-10278.jpg (86051 bytes)

Punta Baja, Spjut 10278,
Mar 1988, Salazinic acid
(TLC Apr 1988)

Bahía de San Quintín,
Spjut, Cassady & McCloud 9327,
May 1986, Salazinic acid
(TLC Nov 1987), isotype

Punta Canoas, Spjut & Marin 11300, Apr 1990, Salazinic acid (TLC Dec 1990).  Growing abundantly on sand.

arenaria-11324.jpg (76488 bytes)

Punta Canoas, Spjut & Marin 11324, Apr 1990, Salazinic acid (TLC Dec 1992)

Punta Canoas
Spjut & Marin 13017,
Apr 1994


Bahía de San Quintín,
Spjut 10241, Mar 1988

Mesa above Punta Rocosa,
Spjut 10365, Mar 1988

Rancho San José between
Puntas Canoas and Blanca, Apr 1990
Spjut & Marin 11395


Mesa southwest of El Rosario,
Spjut 10267, Mar 1988


Near Punta Baja,
Spjut 10272, Mar 1988

Illustration of TLC data showing salazinic acid among other chemotypes of Niebla.

Geographical Distribution


     Niebla arenaria is a fruticose lichen  endemic to Baja California, ranging from Morro Santo Domingo north to near Colonet.  It grows loosely attached to soil on barren sandy beaches along the Pacific Coast and slightly inland on alkaline clay or gravel among low  scrub vegetation, often with Frankenia palmeri and Atriplex julacea, or with Ambrosia spp., Euphorbia misera, and various cacti. 

     Niebla arenaria is distinguished by its chemistry of salazinic acid, which may be accompanied by consalazinic acid, and by its morphology of a hemispherical thallus without a holdfast, intricately divided into numerous narrow ribbon-like branches, appearing antler-like and shortly bifurcate near apex.  Black pycnidia are usually prominent at the tips of shortly bifurcate branchlets.

     Niebla arenaria is often gregarious, presumably from reproducing asexually by  fragmentation of terminal bifurcate branchlets.  It can form a single dominant Niebla community on sandy substrates as observed along the northern shore Bahía Falsa, the type locality, in May 1986, while it also can be member of mixed  Niebla terricolous communities with N. effusa, N. josecuervoi, and N. juncosa on mesas southwest of El Rosario to Punta Baja.  It appears most common in the chaparral-desert transitional region.

     Other species with terminal antler-like branchlets are Niebla limicola and Niebla brachyura. Niebla limicola differs by having flattened and expanded branch parts from which the short bifurcate branchlets develop. It is most common south of Morro Santo Domingo.  Niebla brachyura differs by having hyprotocetraric acid instead of salazinica acid; it  is relatively rare, found mainly in the southern region of the Northern Vizcaíno Desert, and on Isla Cedros.

     The geographical distribution of Niebla arenaria shown on the map of Baja California is based on more than 100 specimens analyzed by thin-layer chromatography (TLC).  The type collection (Spjut, Cassady & McCloud 9327, holotype at US) from Bahía Falsa near San Quintín) included perhaps another 100 thalli that were extracted and tested for anticancer activity.  In addition to isotypes distributed to institutions in Baja California (BCMEX)  and California (LA), other specimens were submitted to the lichen exchange in Arizona (ASU). 

      Although Niebla arenaria can be recognized as described by Spjut (1996), its phylogeny is not resolved (Spjut et al. 2020) along with most species that lack triterpenes while also distinguished by depsidones, except for being absent in N. homaleoides.

     The type for N. arenaria was collected in the vicinity of the type locality for N. pulchribarbara, which was along on a sandy bay shore along Bahía Falsa. The latter differs by having protocetraric acid (Rundel et al. 1972), while the image of the type lacks short bifurcate branching near apex. The type for another related species, N. josecuervoi, which also has salazinic acid, was collected along the same peninsula on a ridge of volcanic rocks. 

     Phylogenetic studies by Spjut et al. (2020) and by Jorna et al. (2021) does not distinguish the species from others in the Depsidone clade, while BPP analysis in Spjut et al. (2020) indicated two species for two collections, 17050 from Bahía de San Quintín, the other from Morro Santo Domingo, 17254. 

Additional References: See Niebla.