Niebla contorta

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

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


W of Baa de Tortugas, towards Punta Eugenia, N 27°49.701, W 115°03.454, 35-40 m, Spjut & Sérusiaux 17178. Jan 2016


W of Bahía de Tortugas, towards Punta Eugenia, N 27°49.701, W 115°03.454, 35-40 m, Spjut & Sérusiaux 17179b. Jan 2016. Occurring with N. spatulata, 17179a (salazinic acid & divaricatic acid).

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Punta Prieta E of Bahía Asunción, BCS
Spjut & Marin 11595

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Punta Eugenia, Spjut & Marin 9733, May 1986

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Arroyo at San Andrés,
Vizcaíno Peninsula,
Spjut 9593, May 1986

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7 mi S of Bahía Asunción, BCS, Spjut 9505. May 1986

7 mi S of Bahía Asunción, BCS, Spjut 9506B (type). May 1986


Population of N. contorta,
near Punta Prieta E of Bah
ía Asunción, BCS 

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Isla Santa Margarita, Spjut & Marin 10692, Apr 1989

Niebla contorta complex-ITS
Group 16



     Niebla contorta is a fruticose lichen endemic to Baja California Sur, occurring mostly in the Southern Vizcaíno Desert and on Isla Santa Margarita in the Magdalena Desert region.  It is distinguished by having divaricatic acid with triterpenes and by the relatively small thallus, <3.5 cm high, with tubular branches near base that become ribbon-like towards apex, appearing flattened, dilated, contorted, undulate, and/or lobulate.  Apothecia when present are submarginal along the apical lobes.

     The type  for Niebla contorta is from the Vizcaíno Peninsula southeast of Bahía Asunción where found on calcareous rocks embedded with fossilized shells. As a species it was observed to form a single dominant community of  Niebla contorta; the thalli appearing fairly uniform in their morphology as shown in top center row of images with reference to Punta Prieta, and by specimens collected on three occasions that were all found to have divaricatic acid with triterpenes.  In the above phylogeny DNA from two type specimens appear in Group 16 represented by two specimens collected within a km of the type locality; the road south from Bahía Asunción appeared to have changed slightly since last traveled there in the 1990's. Another species within several km, N. dilatata, formed a single dominant community on decomposed lava and soil, shown in Group 16 with a difference of one mutation; however, the type locality for N. dilatata is Isla Guadalupe where one might expect to find greater phylogenomic differences. 

     Niebla contorta was rec0gnized to include other morphotypes from different locations on the Vizcaíno Peninsula and on Isla Santa Margarita.  On Punta Eugenia and Punta Cono, the thalli lobes are more tongue-shaped to channeled as in N. dilatata or N. caespitosa.  These similar species differ in having more strongly flattened branches with broader round lobes. Also, the lobes of N. dilatata tend to be thickened along the apical margins, whereas those of N. caespitosa appear thinner towards the margins.  Another related species, N. rugosa, differs by the ladder-like ridging on the upper branches.  Niebla undulata, perhaps most similar, is distinguished by its larger size and often by dark pigmentation spots on the cortex; its type is from southern NVD on a ridge just northwest of Puerto San Andrés.  DNA from a specimen collected in the SVD northwest of Bahía Tortugas, identified N. undulata, however, was found to be within a clade of N. contorta (Spjut et et al. 2020, Fig. 7). 

     The phylogenetic tree shown above is a snip from a larger image of a draft ITS Niebla tree generated May 11, 2016 by Professor Emmanuël Sérusiaux in regard to collections we made in Jan-Feb 2016 and in reference to the 2020 publication cited above (Spjut et al. 2020).   The green, yellow and red colors were added to the lineage-lines by Spjut to distinguish the geographical regions, green for the Chaparral Desert Transition (CDT), red for the Northern Vizcaíno Desert (NVD) and yellow for the Southern Vizcaíno Desert (SVD). Species names and collection numbers were also added by Spjut. The published results using 6-loci (Spjut et al. 2020, Fig. 7) are similar to the ITS except that N. contorta in the ITS tree occurs in two sister clades in which N. contorta  lies within the larger sister clade of N. nr. undulata. Although the 6-loci of Spjut et al. (2020) would seem to provide a finer species resolution, this did not distinguish their occurrence in the CDT region separate from that of the NVD.  Political boundaries were decidedly more practical.  When  phylogeographic  distinction is made based on the ITS phylogenetic tree,  N. contorta appears to have a closer, but disjunct, phylogeographic relationship to the CDT specimens instead of to those in the neighboring NVD region. Other examples are mentioned for vascular plants and for species in Vermilacinia in the Supplement data (S1) file. 

      Additionally, specimens in Group 15 that belong to the green group (N. nr undulata) were collected near El Rosario, whereas those in the yellow sister clade are from near Cerro Elephante (17161) and Bahía Tortugas (17180). They appear to represent cryptic N. contorta from two widely separate locations on the Vizcaíno Peninsula by the topotypes for N. contorta appearing in Group 16, 17140 and 17142.  The other specimens in a separate lineage within this group, 17177 and 17178, are suspected to have perlatolic acid, nevertheless, identified N. contorta (with divaricatic acid); they were collected near Bahía Tortugas. Then there is 17163B—in the N. contorta type Group 16—that was collected near Cerro Elephante on the same rock with 17163A that was found to belong to a separate monophyletic Group—17, sister to Group 16.  Group 18 is similar to a sister subclade in Fig. 7 of Spjut et al. (2020) with N. contorta; their phylogeographical relationships  are to specimens in all three Baja desert regions as shown above but with less agreement among the species identifications; however, the appearance of N. flagelliforma in the phylogenetic tree within this group has similarities to that shown in the phylogenetic trees of Jorna et al . (2021).

For more discussion and reference materials see Introduction to Niebla