Niebla flagelliforma

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

See also Niebla for phylogeography of the genus



South of El Rosario along road to Punta Baja, on sandy, wind-swept ridgeline, Leavitt et al. 16-1019, Dec 2016

Spjut & Sérusiaux 17083A & B, SW of El Rosario, road to Punta Baja, N 30°00.075, W 115°45.965', 140 m Two thalli from the same packet in different ITS clades. Photo editing, auto-contrast the TLC image to accentuate triterpenes, weak in 17083, normal color for divaricatic acid yellow. Note 17080A in ITS phylogeny, N. turgida, in two different clades for two different extracts taken from the same thallus. Lineages colored to indicate Chaparral Desert Transition (green), Northern Vizcaíno Desert (red) and Southern Vizcaíno Desert (yellow. SVD). N. flagelliforma closely related to thalli from the SVD.

Southwest of San Quintín in the Punta Mazo Reserve, on volcanic slopes of Volcan Sudoeste, 5 m
Leavitt et al. 16-731, Dec 2016


Morro Santo Domingo, 130 m
Spjut & Sérusiaux 17273
Jan 2016

Southwest of San Quintín in the Punta Mazo Reserve, on volcanic slopes of Volcan Sudoeste, 5 m
Leavitt et al. 16-856, Dec 2016

Southwest of San Quintín in the Punta Mazo Reserve, west facing slope of Monte Cenizo, 80 m.
Leavitt et al. 16-799, Dec 2016
with a few branches of N. homalea

flagelliforma.jpg (151939 bytes)

San José Ranch between
Punta Canoas and Punta Blanca, Spjut & Marin 11409, Apr 1990

flagelliforma-11240.jpg (26441 bytes)

Punta Canoas, Spjut 11240, Apr 1990

flagelliforma-11401.jpg (80615 bytes)

San José Ranch between
Punta Canoas and Punta
Blanca, Spjut & Marin 11401,
Apr 1990

Mesa Camacho, between
Puerta Catarina and Punta
 Canoas, Spjut & Marin 13065, Apr 1994

Mesa Camacho, between Puerta Catarina and Punta Canoas, Spjut & Marin 13104, Apr 1994

Mesa Camacho, between
Puerta Catarina and Punta
 Canoas, Spjut & Marin 13105,
Apr 1994

Hwy 1 N of Rosarito,
~6 mi S of Punta Prieta
Spjut & Marin 9058A
May 1985

Bahía Santa María,
Spjut & Marin 11383
Apr 1990

Arroyo Sauces, just south of Punta Cuchillo, between Punta Blanca and Punta Canoas
Spjut & Marin 11445, Apr 1990

     Niebla flagelliforma is a fruticose lichen that occurs frequently on wind-sheltered rocks in the Northern Vizcaíno Desert of Baja California, mostly between Guerrero Negro and San Quintín, but possibly further north based on one specimen collected by Palmer from “near San Diego.”  It is recognized by having divaricatic acid (with triterpenes), and a closely reticulate cortex on terminal flagelliform branchlets that are usually densely covered with pycnidia. 

     The flagelliform branchlet appears to be a specialized type of reproductive branch in producing abundant black pycnidia on its cortical ridges in contrast to fewer pycnidia developing on the lower part of the branch, thus, the flagelliform branchlet probably breaks off easily from below—where the main branch widens—which is also where the cortex increases in thickness.  Thalli with fewer flagelliform branchlets generally have fully developed apothecia.  Branchlets that produce apothecia usually change direction of growth at the juncture of apothecia, often nearly at right angles.  Branchlets in some thalli have apothecia that apparently abort development, appearing mostly aggregate near tips of  flagelliform branchlets. 

     Thalli of N. flagelliforma are frequently blackened, appearing as if burnt.  The black areas are associated with an epiphytic cyanobacteria species of Nostoc.  This has also been observed in N. caespitosa and N. podetiaforma.

     Niebla flagelliforma is most likely to be confused with N. caespitosa where they occur together as near Punta Santa Rosalillita. Both species have primary branches that widen above base, but taxonomic emphasis is then placed more on the regular narrow shape of terminal branchlets for identifying N. flagelliforma.

     Other species that resemble Niebla flagelliforma are N. marinii and N. flabellata. They have salazinic acid and pycnidia at the tips of branchlets as well as on lower cortical ridges.  A distinct morph of N. testudinaria, collected by Charis Bratt from the Santa Ynez Mts. and Santa Cruz Is., is similar to N. flagelliforma in the narrow branches with close reticulating cortical ridges, but the branchlets also differ in that they are generally geniculate (bend abruptly), not densely covered with pycnidia, and are shortly bifurcate near apex. 

     Niebla flagelliforma was especially common on rocks of steep slopes east of BCN Hwy. 1 near junction with road to El Tomatal; Spjut & Marin 9058 represents a  voucher for a 100 g sample collected for antitumor screening from this area.

     Two other species with flagelliform branches are noteworthy, N. usneoides (sekikaic acid) and N. isidiaescens (divaricatic acid). Instead of pycnidia, they produce isidia on flagelliform branchlets.

     Niebla flagelliforma phenotypically intergrades with N. homalea and N. juncosa in the Chaparral Desert Transition just north of El Rosario and northern Isla Cedros.

     The phylogeny of Niebla flagelliforma is not only unresolved but puzzling.  17083 collected along the road to Punta Baja from El Rosario was separated into A, having erect branches arising from holdfast, and B, erect branches arising from a prostrate branch. TLC from both specimens were found with divaricatic acid, as might be expected, but weak lacking in triterpenes, compared with other divaricatic species from the same location. Collections by Leavitt et al. also in different clades, e.g., 16-1019 and 16-731. 

     The type is from coastal ridges north of Puerto San Andrés and south of Punta Negra, Spjut 9970.  This site is not easily accessible   Spjut and Sérusiaux collected specimens near Punta Santa Rosalillita; however, no DNA obtained.

Additional References: See Niebla.