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,
Aug 2022

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


Pacific Coast: Punta Morro Santo Domingo, ~40 km N of Guererro Negro; 28°15.771, 114°06.230, 110-130 m. Northern Vizcaíno Desert, Coastal desert shrubs on dunes and among volcanic rocks. One thallus loose on sand, 13 cm diam. Cited in MycoKeys 73: 18, Fig. 7 (2020), Suppl. Data File 3. DNA 4837. TLC: Salazinic acid. Spjut & Sérusiaux 17254, 31 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, ~ 500 m from Type Locality. Thallus intricately divided into narrow segments, not regularly bifurcate, tips with black pycnidia. MycoKeys 73: 18, Fig. 7(2020), Suppl. Data File 3. DNA 4692. TLC: Salazinic acid, protocetraric acid-trace?. Spjut & Sérusiaux 17050-4692, 23 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 its southernmost occurrences near Morro Santo Domingo north to near Colonet (Spjut 1996). It often occurs loosely attached to soil on barren sandy beaches along the Pacific Coast extending 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, or trace of protocetraric 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.  Thalli collected near Punta Baja have longer terminal branches abruptly bifurcated near apex.

     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. Thousands if not millions of thalli are recalled to have covered the shoreline, the cover measured by Rundel et al. (1972) at 70%. However no thalli were seen at type locality in January 2016, although some were encountered ~ 500 m south on less disturbed beaches. 

     Niebla arenaria can also be found in 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 salazinic acid; it  is relatively infrequent, 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 that are segregated by their depsidones (absent in N. homaleoides), and by their by branch morphology (flat vs. tubular-prismatic, irregularly widened vs. sublinear) and their terminal branching (antler-like or terminally dilated and fringed).  Two accessions in Spjut et al. (2020, Fig. 7) are shown in different clades; both Stacey and BPP analyses suggest they represent two species, one collected approximately 500 m from type locality in the Chaparral-Desert Transition near San Quintín, the other from the southernmost region of the Northern Vizcaíno Desert, the type locality for N. marinii.

     Additionally, the type for N. arenaria was collected in the vicinity of the type locality for N. pulchribarbara, a sandy bay shore along Bahía Falsa. The latter distinguished by having protocetraric acid (Rundel et al. 1972; Spjut 1996), and by the lack of 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.  Its branches are tubular-prismatic appearing mostly linear throughout, while often bearing short comb-like branchlets along primary branches; the comb-like branches often much longer than the terminal bifurcations near apex, in contrast to just the relatively short bifurcate branchlets in N. arenaria and N. limicola.

     Additional References: See Niebla.