Subgenus Cylindricaria

©The World Botanical Associates Web Page created June 2017
Content prepared by Richard W. Spjut under webpage subgenus Vermilacinia, April 2003, 
 Subgenus Vermilacinia Webpage revised Sep 2012
Image extracted from Follmann (1966), Nov 2014
Image from Wikipedia added for V. acicularis, Jan 2015
Additional images obtained from review of specimens collected for molecular
 studies undertaken by
Emmanuel Sérusiaux & Richard  Spjut in Jan-Feb 2016
and by Steven Leavitt et al. in Dec 2016, added to webpage June 2017
All images by Richard Spjut unless otherwise indicated.
Images of specimens selected to show representative species of Cylindricaria.
Taxonomy according to the following publications. Last updated Nov 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.
Review at 

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. 2020 Oct 30;74:109-110]

Table of Contents

   Key to the genera Niebla and Vermilacinia
   Key to Subgenera of Vermilacinia and Comments

   Photos of representative specimens for the species in each Subgenus
         Key to the Species of Subgenus Vermilacinia (webpage 2017)
(update with new species:
MycoKeys. 2020 Oct 30;74:109-110]
         Key to the Species of Subgenus Cylindricaria

   Review of Traditional Taxonomy for Cylindricatia.

   Phylogeny of Corticolous Species of Cylindricaria in North America

   Nomenclature (authorities, synonyms, and basionyms)

Key to Niebla and Vermilacinia

Medulla with string-like (chondroid) strands; cortex of 2–3 layers;
      primary branches at least initially with two margins defined by
      cortical ridges that connect and divide at frequent intervals to apex,
      the basal branches bifacial and subtubular (subterete in x-section),
      alternating 90 degrees in plane at various intervals, occasionally
      with prominent inter-marginal secondary
± transverse ridges, or
      secondary transverse ridges common, the  branches then appearing
      prismatic in x-section, or primary branches mostly expanded above
      base, appearing flabellate; zeorin and
[-]-16 α-hydroxykaurane
......................................................................................................... Niebla
Medulla lacking chondroid strands, either with scattered 'knots' of hyphae,
      or of a single cord; cortex  a single layer, either a granular sheath
      or firm crust; primary branches tubular or subtubular with margins
      defined more by facial transverse ridges, the cortex often depressed
      between transverse ridges, marginal ridges when evident not uniting
      and dividing with branch faces alternating 90 degrees; zeorin and/or
[-]-16 α-hydroxykaurane present................................................. Vermilacinia


     Vermilacinia is most easily and definitively distinguished by its secondary metabolites, in which their determination are also needed for identifying the species.  With experience, patterns of surface ridging can be recognized for each of the two genera as outlined in the above key.  Difficulties are encountered with some forms of N. flagelliforma and N. rugosa. Their close reticulate ridging appears much like saxicolous Vermilacinia paleoderma, which often has marginal ridges, but its marginal ridges do not separate and unite at regular intervals. Flabellate species of Vermilacinia, V. varicosa and V. rosei, may be initially seen as Niebla since their branch morphology is rare in Vermilacinia.  When chondroid strands are present, they can usually be seen with the naked eye, but when absent, one never is really sure until a thallus branch is sectioned and viewed under the microscope. 


Key to Subgenera of Vermilacinia

    Cortex crustaceous (like a pie crust), with minute cracks or large
       folds (plicate) that crack, usually >50 μm thick; apothecia lateral
       (facing sideways or perpendicular to branch) or terminal, subterminal
       in one rare species near San Quintín, Baja California; saxicolous or
Subgenus Vermilacinia

    Cortex membranous, 15-30 μm thick; usually with a regular network
      of areolate depressions; apothecia mostly subterminal and appendiculate;
      corticolous or terricolous in South America, corticolous in North America.... Subgenus Cylindricaria


     Subterminal apothecia are generally characteristic of subgenus Cylindricaria, the apothecium usually accompanied by a relatively short continuous branch or spur shoot that ascends abruptly upwards as shown above for V. leopardina.  This is in contrast to subgenus Vermilacinia with terminally positioned apothecia in most North American species, or lateral to subterminal positioned apothecia in South American  species. 

     A mildew-like appearance develops on herbarium specimens of Vermilacinia in approximately six months after a specimen has been collected, and the conditions continues to worsen as time passes. This is the result of efflorescence—a chemical change associated with the breakdown of the cortex through which the internal (medullary) hyphae seem to grow out of; however, efflorescence can be prevented by storing specimens in a refrigerator below 40°F, although there may be a trade-off in that long term storage in a frost-free refrigerator causes the cortex to become brittle over time (observed for ~9 yrs).  The decorticating condition has been attributed to [-]-16 α-hydroxykaurane, a diterpene found in all species of subgenus Vermilacinia, most North American species of subgenus Cylindricaria, but not in Niebla (Spjut 1995, 1996).  This diterpene, also referred to as ceruchinol, is apparently rare in lichens, although known in mosses (Physcomitrium).   Efflorescence is common in most species of subgenus Vermilacinia  and  in some species of Cylindricaria that contain bourgeanic acid or the T3 compound (Spjut 1996); the Cylindricaria species include the  sorediate V. cephalota, V. cerebra, V. leonis, and the chemically related nonsorediate V. cerebra and V. tigrina.  The terpenoid compounds that characterize Vermilacinia have not been reported in Niebla (Spjut 1996).  Indeed, Niebla and Vermilacinia have little in common in their major secondary metabolites. 

     The occurrence of ceruchinol in Vermilacinia appears closely associated with bourgeanic acid, but only in the New World.  A ‘bourgeana group’ within the genus Ramalina is recognized for  Macaronesian and Mediterranean species of Ramalina (Sérusiaux et al. 2010; Spjut et al. 2020). This was treated in the genus Niebla by Rundel and Bowler (1978) and Bowler & Marsh (2004), based also on medullary chondroid strands that are interconnected across the medulla, but they are not isolated from the cortex except for the rare R. portosantana, which differs by producing pseudocyphellae, and has been found more related to species within the ‘decipiens group’ (Spjut et al. 2020).



Images below include 2016 collections pertaining to field work and studies on

Evolutionary history of coastal species of fog lichen genera
Niebla, Ramalina and Vermilacinia

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

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


V. cephalota
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17249b, Feb 2016

V. cephalota
West of Villa Jesus Maria, along shoreline, at Punta Morro Santo Domingo, 10 m, Leavitt et al. 16-922, Dec 2016

V. cephalota
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17248c, Feb 2016

V. cephalota
West of Villa Jesus Maria, along shoreline, at Punta Morro Santo Domingo, 19 m, Leavitt et al. 16-921, Dec 2016

V. cephalota
Laguna and peninsula of San Quintín, N 30°27.442' W, 116°00.794', 20 m, Spjut & Sérusiaux 17040, Jan 2016

cephalota9718.jpg (81725 bytes)

V. cephalota
Sierra Hornitos, Vizcaíno Peninsula, Spjut 9718, May 1986


cephalota.jpg (83386 bytes)

V. cephalota
Between Punta Rocosa
and Punta Negra, BCN
Spjut & Marin 9076, May 1985


V. cerebra
Laguna and peninsula of San Quintín, N 30°29'743, W 116°00.044', 4 m, Spjut & Sérusiaux 17033, Jan 2016

V. cerebra
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17266a, Feb 2016

V. cerebra
Vicinity of Punta Catarina, south of point, on gypsum-based badlands, 50 m, Leavitt et al. 16-1023, Dec 2016

V. cerebra
Punta Canoas, on gravel and small rocks, in a region dominated by alkaline soils, 65 m, Leavitt et al. 16-956, Dec 2016

V. cerebra
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17249d, Feb 2016

V. cerebra
Southwest of San Quintín in the Punta Mazo Reserve, volcanic slopes of Volcan Sudoeste, 40 m, Leavitt et al. 16-921, Dec 2016



cerebra-9722.jpg (103565 bytes)

V. cerebra
Near Rosarito, BCN
Spjut 9722, May 1986


cerebra-9796.jpg (128477 bytes)

V. cerebra
Morro Santo Domingo, BCN
Spjut 9796, May 1986

V. cerebra
Just S of El Rosario, BCN
Spjut 10253, Mar 1988
with zeorin, hydroxykaurane, salazinic acid

V. cerebra
Punta Rocosa, BCN
Spjut 10387, Mar 1988
zeorin, hydroxykaurane, 3 unknowns, bourgeanic acid, usnic acid

V. cerebra
May 1986, Vizcaíno Peninsula

V. corrugata
Laguna and peninsula of San Quintín, N 30°29'743, W 116°00.044', 4 m, Spjut & Sérusiaux 17031, Jan 2016 , zeorin, ± usnic acid


V. corrugata

SE of El Rosario along Hwy 1, N 30°02.833 W, 115°33.734, 230 m, Spjut & Sérusiaux 17095, Jan 2016,  Zeorin, ± T4, ± atranorin

V. corrugata

 Ca. 15 km, N of Guerrero Negro,  N 28°12.575, W 113°51.664, 55 m,  Spjut & Sérusiaux 17240, Feb 2016, zeorin, T3, usnic acid

V. corrugata
Laguna and peninsula of San Quintín, N 30°29'743, W 116°00.044', 4 m, Spjut & Sérusiaux 17025, Jan 2016 , zeorin, ± usnic acid

V. corrugata
Vizcaíno Peninsula. Along the road from Bahía de Tortugas to Vizcaíno, N 27°37.826, W 113°25.190, 70 m, Spjut & Sérusiaux 17236a, Jan 2016, zeorin


V. corrugata
Along shoreline, Punta Morro Santo Domingo, 10 m, Leavitt et al. 16-916, Dec 2016

corrugata-10723.jpg (244788 bytes)

V. corrugata
Inland from Puerto Cancun,
BCS, Spjut 10723

V. flaccescens
Chile, Juan Fenandez Island
Tucker Herb. (FH)

V. flaccescens
Chile, Papudo Prov.
Looser, Dec. 1928
Tucker Herb. (FH)

V. howei
Vizcaíno Peninsula, between Bahía de Tortugas and Punta Eugenia,  N 27°44.969, W 114°56.690 140-160 m, Spjut & Sérusiaux 17195(B), Jan 2016

V. howei
Vizcaíno Peninsula, between Bahía de Tortugas and Punta Eugenia,  N 27°44.969, W 114°56.690 140-160 m, Spjut & Sérusiaux 17189, Jan 2016, zeorin

V. howei
Vizcaíno Peninsula, between Bahía de Tortugas and Punta Eugenia,  N 27°44.969, W 114°56.690 140-160 m, Spjut & Sérusiaux 17187, Jan 2016, zeorin, ± T3

V. howei
Vizcaíno Peninsula, between Bahía de Tortugas and Punta Eugenia, N 27°49.701 W, 115°03.454, 35-40 m, Spjut & Sérusiaux 17172, Jan 2016

V. howei
Vizcaíno Peninsula, between Bahía de Tortugas and Punta Eugenia, N 27°49.701 W, 115°03.454, 35-40 m, Spjut & Sérusiaux 17171, Jan 2016

vermilacinia_howei_9722.jpg (84534 bytes)

V. howei
Vizcaíno Peninsula,
Sierra San José de Castro
Spjut 9722, May 1986

vermilacinia_howei_9686.jpg (45718 bytes)

V. howei
Vizcaíno Peninsula,
hills N of Bahía de Tortugas
Spjut 9686, May 1986


vermilacinia_howei_9713.jpg (118336 bytes)

V. howei
Vizcaíno Peninsula,
Sierra Hornitos
Spjut 9713, May 1986


leonis-12692.jpg (185568 bytes)

V. leonis
Inland from Puerto Cancun,
BCS, Spjut 12692

V. leonis
Chile, Santiago
Looser 838 (FH)


leonis-9606D.jpg (117478 bytes)

V. leonis
Sierra Hornitos,
Vizcaíno Peninsula, 
Spjut 9606, May 1986


V. leopardina
NE of San Rosalillita, narrow valley in the mesa, N 28°40.572, W 114°13.736, 35 m, Spjut & Sérusiaux 17310, Feb 2016

V. leopardina
Along shoreline, Punta Morro Santo Domingo, 10 m, Leavitt et al. 16-916, Dec 2016


V. leopardina
Laguna and peninsula of San Quintín, N 30°26.988' W 115°59.659' 1 m, Spjut & Sérusiaux 17038, Jan 2016

V. leopardina
West of Transpeninsular Highway 1, along road to El Consuelo, west of Canada de San Quintín, in coastal dunes, 12 m, Leavitt et al. 16-1073




V. leopardina
Vicinity of Punta Catarina, south of point, on gypsum-based badlands, 50 m, Leavitt et al. 16-1036, Dec 2016


V. leopardina

Vizcaíno Peninsula. Along the road from Bahía de Tortugas to Vizcaíno, N 27°37.826, W 113°25.190, 70 m, Spjut & Sérusiaux 17236c,e, Jan 2016, zeorin




leopardina-9986-isotype.jpg (66912 bytes)

V. leopardina
Ridge S of Punta Negra, BCN
Spjut 9886, isotype (wba)


leopardina9334.jpg (82766 bytes)

V. leopardina
Bahía de San Quintín
Spjut 9334, May 1986

V. nylanderi
E of Bahía de Ascunci
ón, near the coast, N 27°09.813, W 114°14.752, 20 m, type locality, Spjut & Sérusiaux 17137, Jan 2016

V. nylanderi
W of Vizcaíno, along the road to Punta Eugenia, N 27°30.841, W 114°19.167, 235 m, Spjut & Sérusiaux 17211, Jan 2016

V. nylanderi
W of Vizacaíno, along the road to Punta Eugenia, N 27°30.841, W 114°19.167, 235 m, Spjut & Sérusiaux 17211, Jan 2016

V. nylanderi
S of Punta Prieta, along Hwy 1, N 28°49.377, W 114°06.586, 210 m, Spjut & Sérusiaux 17099, Jan 2016


V. nylanderi
Along gravel to Punta Catarina, ca. 2 km inland from coast, on silty flat, growing on euphorbs, Leavitt et al. 16-1063

vermilacinia_nylanderi_9336.jpg (77237 bytes)

V. nylanderi
E of El Rosario,
along Hwy 1, 
Spjut 9336, May 1986

vermilacinia_nylanderi_9536.jpg (54524 bytes)

V. nylanderi
Vizcaíno Peninsula,
~7 mi S of Bahía Asunción
Spjut 9536A, May 1986

vermilacinia_nylanderi_1159.jpg (81977 bytes)

V. nylanderi
Vizcaíno Peninsula,
Punta Prieta W San Hipolito,
E of Bahía de Asunción
Spjut & Main 11598, Apr 1990

V. tigrina
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17253, Feb 2016, zeorin, [-]-16 α-hydroxykaurane, salazinic acid

V. tigrina
South of El Rosario along road to Punta Baja, on sandy, wind-swept ridgeline, 80 m, Leavitt et al. 16-1010, zeorin, [-]-16 α-hydroxykaurane, ±T3, salazinic acid, traces of 3 unknowns,

V. tigrina
West of Villa Jesus Maria, along shoreline, at Punta Morro Santo Domingo, 10 m, Leavitt et al. 16-914, Dec 2016, zeorin, [-]-16 α-hydroxykaurane, traces of 2 unknowns, salazinic acid, consalazinic acid

vermilacinia_tigrina_10601.jpg (239789 bytes)

V. tigrina
1–2 miles
W of Puerto Cancun, BCS
Spjut & Marin 10602, Apr 1990

V. tigrina
1–2 miles
W of Puerto Cancun, BCS
Spjut & Marin 10606, Apr 1990

Illustration of TLC Data
for species of Vermilacinia.
Colors shown uner UV long wavelength after plate
has been charred.

V. tigrina
Follmann (1966, Ramalina tigrina), Fig. 4:
Chile. Atacama Desert:
Antofagasta, Cerro Moreno, 600 m

V. zebrina
Holotype, from near Arcata, California

V. zebrina
N of Guerrero Negro, Punta Santo Domingo, N 28°15.269, W 114°06.378, 110-130 m, Spjut & Sérusiaux 17253, Feb 2016, zeorin, [-]-16 α-hydroxykaurane, salazinic acid, occurring with V. tigrina

Laguna and peninsula of San Quintin, underside of rock overhang, 40 m, Spjut & Sérusiaux 17071 reassigned to 17068, Jan 2016




 Subgenus Cylindricaria


Review of Traditional Taxonomy

Phylogeny of Corticolous Species in North America


Review of Traditional Taxonomy

     Vermilacinia Subgenus Cylindricaria is easily recognized by the thallus divided into cylindrical branches from a holdfast attached to bark of shrubs or trees, rarely on rock as reported for sorediate species V. cephalota and V. tigrina. The cortex is membranous and relatively thin (15–30 µm) over a loose network of longitudinally oriented medullary hyphae that converge at regular intervals into  knotted bundles. This network arrangement allows for the collapsing and expansion of the cortical matrix onto the cortical ridges as desiccation (dissipation) and hydration develops from the advance and dissipation of fog.  Subgenus Cylindricaria is further characterized by having black pycnidia that evidently develop into larger black spots or transverse bands, and by the absence of distinct medullary chondroid strands, and by apothecia spur shoots (appendiculate).

         It should be noted that the name "Niebla ceruchis," regarded as a synonym of V. ceruchis (Spjut 1996), applies to a terricolous and saxicolous species in South America in subgenus Vermilacinia. This is based on the type specimen collected by Menzies—described and named by Acharius (1803) Parmelia ceruchis.  The lectotype designated by Spjut (1996) appears to be from a terricolous thallus judging from its morphological and chemical attributes similar to specimens at Harvard (FH) reportedly  collected by Darwin from the coastal South American desert in northern Chile. Darwin noted that it was the only “plant” lying loose on sand for “16 leagues inward.”   The Darwin specimens studied in detail by Spjut (1996) were found to have a relatively thick firm cortex characteristic of subgenus Vermilacinia.  TLC of a Darwin specimen revealed three terpenoid compounds as seen in an image of the TLC data for the type of V. ceruchis.  Thus, V. ceruchis is clearly more related to the saxicolous V. combeoides than to the corticolous species. Vermilacinia tigrina—not mentioned by Bowler and Marsh (2004) in their revision of Niebla—appears to be the earliest species name for the corticolous group, distinguished by Bowler (1981) and similarly by Spjut (1996) from the saxicolous species group. 

     Notwithstanding, the name Niebla ceruchis has continued to be misapplied to corticolous species in North America (Bowler & Marsh 2004). The application of this name to North American corticolous species apparently originated from Howe (1913), although Tuckerman (1866) had earlier reported that he had received specimens identified as Ramalina ceruchis.  They were collected on shrubs near San Diego and San Francisco, but this was before Nylander's (1870) monograph.  Nylander (1870) considered the North American corticolous species to belong to what is now a saxicolous species of Niebla (Ramalina) testudinaria. Evidently, the honeycomb-like cortical surface seen in N. testudinaria and V. leopardina was considered by Nylander to be diagnostic  for treating the variation under the same species (Ramalina testudinaria).  Ramalina ceruchis was considered by Nylander (1870) to occur only in South America, on trees and shrubs in Chile and Peru, and his description of a new North American species, R. combeoides, was placed closer to R. ceruchis than to R. testudinaria

    Again, key differences between the two subgenera were recognized by Bowler (1981) based on his detailed anatomical study of the cortex of many Ramalinacea genera, but the wrong species name was applied to the corticolous Niebla group. This may be due in part to not having studied all the relevant type specimens of Ramalina. As reported by Spjut (1996), species indicated to have specific lichen substances lacked reference to type specimens; for example, the depside tumidulin (= 3,5 methy dichlorolecanorate) reported by Follmann (1966) to have been found in  Ramalina tumidula, treated as a synonym of V. ceruchis by Spjut (1996), was not present in the type specimen. The misapplication of names are applied in later studies, leading to further confusion as to which lichen substances may be present in a species. For example, specimens of subgenus Vermilacinia in the U.S. Natl. Herbarium, had been erroneously annotated by Rundel and Bowler—in their study of Niebla—to have methyl 3,5 dichlorolecanorate.  Specimens of Vermilacinia cedrosensis and V. paleoderma collected by Spjut in Baja California and analyzed by chemists of the Natural Products Group in the National Cancer Institute could not identify any methy dichlorolecanorate. Subsequently, Spjut (through Mason Hale) sent a South American specimen of subgenus  Cylindricaria—that he tentatively identified as having the depside—to C. Culberson who—along with A. Johnson–confirmed its presence. This depside  was then recognized by Spjut (1996) as a key taxonomic character for V. flaccescens, a corticolous species he regarded endemic to South America.  More recently,  it seems that  the triterpene zeorin was confused with the diterpene in V. tuberculata (Niebla t., Riefner et al. 1995; Spjut annotation labels for COLO loan). 

     Two common species, Vermilacinia leopardina and V. corrugata, have been recognized by their cortical differences. They appear related to zonal moisture differences correlated with the occurrence of the secondary metabolite diterpene [-]-16 α-hydroxykaurane.  For example, in the Northern Vizcaíno Desert of Baja California, V. leopardina, which has the diterpene, is more common along the immediate coast.  In contrast, V. corrugata, which lacks the diterpene, occurs further inland, or on leeward slopes where moisture from fog dissipates.  The cortical surface of the former species generally appears smooth in texture, whereas in the latter, the cortex is deeply and/or sharply rugose.  On the Vizcaíno Peninsula (Southern Vizcaíno Desert), a reverse relationship is evident.  The absence of the diterpene is seen in thalli with a relatively smooth cortical features, usually with crater-like rounded depressions; they are referred to V. howei. which occurs more often near the coast than V. nylanderi, a species with a strongly rugose cortical surface that contains the diterpene. Vermilacinia howei appears endemic to northwestern  Vizcaíno Peninsula, while the rare occurrences reported on the main peninsula of Baja California and in the Channel Islands differ by shorter thallus with erect capillary branches lacking apothecia, and may prove to a distinct species.   

    Another feature of taxonomic significance in Cylindricaria is the development of soralia.  At least four species have soralia (only one species recognized in Bowler & Marsh 2004).  The sorediate species, and two related nonsorediate species, V. cerebra, and V. tigrina, produce bourgeanic acid and depsidones.  Additionally, Sipman (2011) recognized a South American species with apical soralia, Niebla granulans, in contrast to lateral soralia seen in the other species, although in some V. cephalota they appear restricted terminally on short pinnately arranged branchlets for a short distance near the end of a primary branch.  This was transferred to Vermilacinia by Spjut and Sérusiaux  (Spjut et al. 2020, V. granulans). Also, V. cerebra may appear sorediate terminally by aborted development of apothecia where the cortex appears to slough off  with age.  Vermilacinia zebrina, recognized by Spjut (1996), differs from V. cephalota  in its much dichotomously branched thallus of capillary branches, the branching more frequent towards apex; the interval branches thus becoming shorter in length.  This species seems most common in the Channel Islands, while it also occurs in Namibia (image in V. Wirth, Lichens of the Namib Desert, Hess, Göttingen. 2010). Intermediates appear evident by the  spiculiferous branchlets developing nearer the apex of  primary branches. These intermediates may be separated by the presence/absence of depsidones (Spjut 1996); however, thalli with salazinic acid collected in Baja California on rocks or shrubs  appear to belong to V. zebrina by the relatively uniform shape of the branches from the thallus base. Finally, V. leonis, a species found  in BCS and in South America, is distinguished by a thallus of interwoven capillary flexuous branches. In BCS it commonly occurs with V. tigrina and species of Ramalina.

     The South American species of Cylindricaria include chemotypes not found in North America. Vermilacinia flaccescens, as mentioned above, was broadly interpreted to include any Ramalina thallus with  methyl 3,5 dichlorolecanorate.  The depside has also been reported in South American species of Ramalina (Spjut 1996).  This may have been acquired from hybridization (or via horizontal gene transfer). Similar thalli with this depside but without the terpenes  have been referred to Vermilacinia cactacearum (Follmann) Follmann & Werner (Schedae ad Lichenes Exsiccati).

     Another South American species, Niebla nashii,  reportedly has tumidulin and terpenes; however, the terpenid compounds were not clearly specified. Its placement in the genus Niebla or Vermilacinia is therefore questionable, especially since it lacks pycnidia as well as apothecia (Sipman 2011).  It was also described to have medullary chondroid strands free from the cortex, while its overall resemblance is to Ramalina lacera (With.) J. R. Laundon.  Additionally, the species reportedly has more than one depside, suggesting affinities to the Old World Bourgeana clade of Ramalina (Krog, H. & H. Østhagen. 1980; Sérusiaux et al. 2010).  Moreover, "Vermilacinia lacera" (With.) Follm. & Wern. was proposed (G. Follmann & B. C. Werner: Sched. Lich. Exs. Univ. Coloniensis Ed., Fasc. 24, 11. 2003. No. 479), evidently based on chemotaxonomy.

     The evolutionary ties of North American subgenus Cylindricaria to its South American species are evident in the sorediate species (V. leonis) and the non-sorediate V. cerebra and V. leopardina.  The sorediate species and V. cerebra, which were recognized to occur in both Americas, share other key metabolites such as depsidones and bourgeanic acid.   Vermilacinia tigrina thalli in South America include a corticolous form with hypoprotocetraric acid and a terricolous form with psoromic acid. In North America, it is corticolous with either norstictic acid, or salazinic acid, or rarely both depsidones are present.   Vermilacinia leopardina, distinguished by lacking depsidones, has the same secondary metabolites in both Americas, mostly zeorin and [-]-16 α-hydroxykaurane.  The North American thalli referred to V. tigrina, which often differ from V. leopardina in having longer branches with a more sharply corrugated cortex. may prove to be a distinct species, especially if the South American thalli that show differences in habit and chemistry, justify separate species status.

     The evolutionary trend in Cylindricaria seems to be loss of secondary terpenoid metabolites, which may have occurred after Cylindricaria were established in both North America and South America. This seems supported by phytogeographcal data for the species on each continent.  Examples are the reversed pattern in the diterpene chemistry between the Vizcaíno and northern peninsular species that may have since evolved independently—in the loss of the diterpene—at a time when the Vizcaíno Peninsula was perhaps an island separated from the main Baja peninsula.  However, this interpretation is supported only partially in the DNA phylogenetic study by reference to a black banded species group that lacks salazinic acid and the terpene T3 (Spjut et al. 2020).  Moreover, tumdilin previously thought to occur only in South American corticolous V. flaccescens has been discovered by Spjut et al. (2020) to occur in a newly described saxicolous species of Vermilacinia endemic to Baja California.

Phylogeny of Corticolous Species in North America

      Phylogeny of the species in subgenus Cylindricaria is supported for a V. corrugata group characterized by the absence of black bands and absence of ceruchinol (Spjut et al. 2020). The V. corrugata group comprises several species, whereas Spjut (1996) described only one species.  The type was collected in Baja California  Sur in the vicinity of Ciudad Insurgentes where Spjut & Sérusiaux did not visit in 2016.  These species  probably can be recognized based on >100  herbarium specimens that were segregated by the author (Spjut 1996) after his publication for possible distinction of varieties. 

     The sorediate species, V. cephalota, includes at least two species, one in California where the type was collected and another in Baja California. Vermilacinia  cerebra, generally recognized by terminal swollen branches or with clusters of terminal aborted development of apothecia, is supported.  Vermilacinia leonis described from Baja California Sur was not collected for Spjut et al. (2020).  

     The black-banded species group (V. howei, V. leopardina, V. nylanderi, V. tigrina, V. zebrina), variable in chemistry and morphology, needs more study. Although V. howei is easily distinguished by the absence of ceruchinol, it was not supported in ITS phylogenetic trees, whereas a terpene referred to as T3 defines a group for specimens largely identified V. tigrina and V. cerebra (Leavitt et al. 2016 specimens), that also have salazinic acid.  The type for V. tigrina, distinguished by producing psoromic acid,  was collected in Chile where likely  it will be proved endemic. Vermilacinia nylanderi and V. leopardina, which lack salazinic acid, generally differ by the cortex appearing strongly shriveled and dull in the former, and smoothly dimpled and shiny  in the latter.   DNA phylogeny of these species fall into two sister groups in which V. leopardina occurs both  while V. nylanderi is restricted to one.  Although V. zebrina was not supported, the Baja California specimens differ from the  typical California specimens by the thallus branches of ± uniform cylindrical shape disarticulating at frequent intervals from near base whereas California specimens break apart more near the apex where dichotomously much branched.

Key to Species of Subgenus Cylindricaria 

1(0).     Thallus with methyl 3,5 dichlorolecanorate; South America.
1a With
[-]-16 α-hydroxykaurane.........................................................................1b
1a Without
[-]-16 α-hydroxykaurane................................. Vermilacinia cactacearum
1b. Thallus with terminal punctiform soralia..................... Vermilacinia granulans

                      1b. Thallus not sorediate...............................................Vermilacinia flaccescens
1.         Thallus lacking methyl 3,5 dichlorolecanorate; North and
South America.......................  2

2(1).    Thallus with distinct rounded soralia on acicular to flexuous branchlets
                (best observed in the field)....................................................................................  3
2.         Thallus not sorediate (sometimes appearing sorediate or moldy in the herbarium
                 as a result of chemical sublimation in which the cortex breaks down and
                 whitish crystalline deposits appear along with medullary hyphae that may
                 spread out through the cortical cracks or apertures)................................................. 5

3(2).     Branches usually dilated or inflated, or more than 1 mm wide, rounded (obtuse)
                 to apex on some branches, with or without acicular branchlets; variable in shape;
                 cortex dark green to blackish green, with or without irregular black patches; soralia
                 irregularly round, rash-like................................................... Vermilacinia cephalota
3.         Branches all ± of uniform width, < 1.0 mm wide; acicular near apex; cortex pale in
                 color, yellowish-green or straw-colored; soralia disciform to capitular........................ 4

4(3).     Thallus regularly dichotomously divided, shortly bifurcate near apex, with or without
                  black bands and spots; soralia bluish gray, capitular:
                     4a Thallus with well defined black spots or transverse bands; California........ 
                          .................................................................................... Vermilacinia zebrina
                     4b Thallus without regular black bands or spots; Bahía Asuncion,
                          Vizcaíno Peninsula................................................................ Vermilacinia sp.
Thallus irregularly branched; without regular black spots and bands; soralia
                          disciform, grayish-white; Baja California Sur, South America.Vermilacinia leonis

5(2).    Thallus containing the diterpene [-]-16 α-hydroxykaurane.............................................. 6
5.        Thallus lacking the diterpene [-]-16 α-hydroxykaurane................................................... 9

6(5).     Apothecia appearing to abort development, terminally aggregate on expanded
                  lobes; black bands or spots not well defined; bourgeanic acid present, thallus
                  usually developing whitish mold-like deposits within six months when kept
                  at room temperature; mostly away from the immediate coast.....Vermilacinia cerebra

6.        Apothecia usually subtended by a capillary branchlet; black bands often regularly
                  present; chemistry variable.................................................................................. 7

7(6).     Thallus with depsidones (medulla PD+; hypoprotocetraric acid, or norstictic acid,
                  or psoromic acid, or salazinic acid in weak to strong concentrations); black
                  band and spots regularly present or absent; a variable species distinguished
                  primarily by the presence of depsidones; mostly Baja California Sur, originally

described from South America............................................... Vermilacinia tigrina
7          Thallus lacking depsidones (medulla PD-).................................................................. 8

8(7).     Cortical surface with irregular shallow depressions, appearing mostly smooth;
                  black transverse bands and/or spots regularly present; common near the
                  immediate coast,
peninsular CA & BCN, Channel Islands, also in South
                  America.......................................................................... Vermilacinia leopardina
        Cortical surface deeply folded and/or regularly lacunose; black
transverse bands
                  or enlarged spots absent or not regularly present; mostly Channel
                  and Vizcaíno Peninsula, also in South America.................... Vermilacinia nylanderi

9(5).     Branches with black bands or elongated black spots; cortex generally smooth
                  with rounded shallow depressions; infrequent, Channel Islands, San

                  Quintín peninsula, and western
Vizcaíno Peninsula....................Vermilacinia howei
9.         Branches lacking black bands or elongated spots, often acutely 5-ridged; cortex
                  acutely lacunose; common, especially near the perimeter of fog zone,
                  Vizcaíno and Magdalena Deserts........................................Vermilacinia corrugata



Acharius, E. 1803. Methodus Qua Omnes Detectos Lichenes. Stockholm

Bowler, P.A. 1981. Cortical diversity in the Ramalinaceae. Canad.
J. Bot. 59:437-453.

 __________, R. E. Riefner, Jr., P. W. Rundel, J. Marsh & T.H.
Nash, III. 1994. New species of Niebla (Ramalinaceae) from
western North America. Phytologia 77: 23-37.

__________ and J. Marsh. 2004. Niebla. ‘Lichen Flora of the Greater Sonoran Desert 2’: 368–380.

Diederich, P.  2007. New or interesting lichenicolous Heterobasidiomycetes. Opuscula Philolichenum 4: 11–22.

Follmann, N. G. (1994). Darwin's “lichen oasis” above Iquique, Atacama Desert rediscovered. International Lichenological Newsletter 27: 23-25.

Herre, A. W. C. T. 1906. The foliaceous and fruticose lichens of the Santa Cruz Peninsula, California. Proc . Acad. Sci. Washington 8: 325–396.

Howe, R.H., Jr. 1913. North American species of the genus Ramalina. The Bryologist 16: 65–74

Krog, H. & H. Østhagen. 1980. The genus Ramalina in the Canary Islands. Norwegian J. Bot. 27:255-296.

Marsh, J. & T.H. Nash, III. 1994.  A new lichen species, Niebla
, is described from Baja California, Mexico. Phytologia
76: 458-460.

Nylander W. 1870. Recognitio monographica Ramalinarum. Bull. Soc. Linn. Normandie, Sr. 2, 4:101–181.

Riefner Jr., R. E., P. A. Bowler, J. Marsh & T. H. Nash III. 1995.
Niebla tuberculata (Ramalinaceae): A new lichen from California. Mycotaxon 54: 397-401.

Rundel, P.W.,  P.A. Bowler & T.W. Mulroy. 1972. A fog-induced
lichen community in northwestern Baja California, with two new species
of Desmazieria. Bryologist 75:501-508.

Rundel, P.W. and  P.A. Bowler, 1978. Niebla, a new generic name
for the lichen genus Desmazieria (Ramalinaceae). Mycotaxon

Sérusiaux, E., P. Van den Boom, and D. Ertz. 2010. A two-gene phylogeny shows the lichen genus Niebla (Lecanorales) is endemic to the New World and does not occur in Macaronesia nor in the Mediterranean basin.  Fungal Biology 114: 528-37.

Sipman, H.J.M. 2011.  New and notable species of Enterographa, Niebla, and Sclerophyton s. lat. from coastal Chile. Bibliotheca Lichenologica 106: 297-308.

Smithsonian Institution. 1990 (Jan. 9). National Museum of Natural History. Letter from the Director to Dr. Richard Spjut indicating renewal of his appointment as Associate and Collaborator, with particular emphasis on Dr. Spjut's study of the lichens of Baja California.

Spjut, R. W. 1995. Vermilacinia (Ramalinaceae, Lecanorales), a new genus of lichens. Pp. 337-351 in Flechten Follmann; Contr. Lichen. in honor of Gerhard Follmann, F. J. A. Daniels, M.
Schulz & J. Peine, eds., Koeltz Scientific Books, Koenigstein.

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_________. 1997. The California Floristic Element on Isla Cedros. Paper presented at the Baja California Botanical Symposium, Aug 14-16, Museum of Natural History, San Diego, Abstract.

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Vermilacinia acicularis Spjut, Sida Botanical Miscellany 14: 152. 1996.
Vermilacinia albicans Spjut ined. = V. cedrosensis
Vermilacinia cactacearum (Follmann, Philippia 3: 1976) Follmann & Werner.  Schedae ad Lichenes
         Exsiccati Selecti ab Instituto Botanico Universitatis Coloniensis Editi XXVIII. Fasciculus
         Numeris 541–560, 2004.  Basionym: Desmazieria cactacearum.
Vermilacinia cedrosensis (Marsh & Mash, Phytologia 76: 459. 1994) Spjut, Sida Botanical Miscellany
         14: 153. 1996.  Basionym: Niebla cedrosensis
Vermilacinia cephalota (Tuckerman, Synop. N. Amer. Lich. 21. 1882)  Spjut & Hale,
         Flechten Follmann 347. 1995.  Basionym: Ramalina ceruchis f. cephalota
Vermilacinia cerebra Spjut, Sida Botanical Miscellany 14: 181, 1996.
Vermilacinia ceruchis (Acharius, Methododus 260. 1803) Spjut & Hale, Flechten Follmann 345. 1995.
          Basionym: Parmelia ceruchis
Vermilacinia ceruchoides (Rundel & Bowler, Phytologia 77: 26. 1994) Spjut, Sida Botanical
         Miscellany 14: 152. 1996.  Basionym: Niebla ceruchoides.
Vermilacinia ceruchoides
Spjut ined. = Vermilacinia acicularis
Vermilacinia convoluta
Spjut ined.
Vermilacinia combeoides (Nylander, Bull. Soc. Linn. Normandie S
ér 2, 4: 107. 1870) Spjut & Hale,
         Flechten Follmann 345. 1995. Basionym: Ramalina combeoides
Vermilacinia corrugata Spjut, Sida Botanical Miscellany 14: 183. 1996.
Vermilacinia flaccescens (Nylander, Bull. Soc. Linn. Normandie S
ér 2, 4: 109. 1870) Spjut & Hale,
         Flechten Follmann 348. 1995.  Basionym: Ramalina flaccescens
Vermilacinia granulans
(Sipman) Spjut & S
Vermilacinia halei
Spjut ined. = Vermilacinia laevigata
Vermilacinia howei
Spjut, Sida Botanical Miscellany 14: 187, 1996.
Vermilacinia johncassadyi Spjut, Sida Botanical Miscellany 14: 162. 1996.
Vermilacinia laevigata (Bowler & Rundel, Phytologia 77: 31. 1994) Spjut, Sida Botanical
         Miscellany 14: 163. 1996.  Basionym: Niebla laevigata
Vermilacinia leonis
Spjut, Sida Botanical Miscellany 14: 189. 1996.
Vermilacinia leopardina Spjut, Sida Botanical Miscellany 14: 190. 1996.
Vermilacinia ligulata Spjut, Sida Botanical Miscellany 14: 165. 1996.
Vermilacinia nylanderi Spjut, Sida Botanical Miscellany 14: 192. 1996.
Vermilacinia paleoderma Spjut, Sida Botanical Miscellany 14: 7, 166. 1996.  Note: Type
         was stated on p. 166 to be Spjut & Marin 9074 from near San Andr
és Ranch, which
         complies with ICBN Art, 37.6, and "holotype" was stated on p.7 to have been deposited in the
         United States National Herbarium (US), in accordance with Art. 37.7. Isotypes were sent
         to BCMEX and LA. Name was validly published.
Vermilacinia polymorpha
(Bowler, Marsh, Nash & Riefner, Phytologia 77: 33. 1994) Spjut, Sida
         Botanical Miscellany 14: 168. 1996.  Basionym: Niebla polymorpha
Vermilacinia procera
(Rundel & Bowler, Phytologia 77: 34. 1994) Spjut, Sida Botanical
         Miscellany 14: 152. 1996.  Basionym: Niebla procera
Vermilacinia pulvinata
Spjut ined. = Vermilacinia ceruchoides
Vermilacinia pumila
Spjut, Sida Botanical Miscellany 14: 169. 1996.
Vermilacinia reptilioderma Spjut, Sida Botanical Miscellany 14: 171. 1996.
Vermilacinia reticulata Spjut & Sérus MycoKeys 73: 1–68. Mycobank
Vermilacinia rigida Spjut, Sida Botanical Miscellany 14: 172. 1996.
Vermilacinia robusta (Howe
, Bryologist 16: 73. 1913) Spjut & Hale,
         Flechten Follmann 348. 1995. Basionym: Ramalina combeoides var. robusta
Vermilacinia robustiella
Spjut ined = Vermilacinia procera
Vermilacinia roccellaeformis (Bory) Spjut ined.
Vermilacinia rosei
Spjut, Sida Botanical Miscellany 14: 175. 1996.
Vermilacinia subterminalis Spjut ined.
Vermilacinia tigrina (
Follmann & Huneck, Willdenowia 6: 208. 1969) Spjut & Hale,
         Flechten Follmann 348. 1995. Basionym: Ramalina tigrina
Vermilacinia tuberculata
(Riefner, Bowler, Marsh & Nash, Mycotaxon 54: 397. 1995) Spjut
         Sida Botanical Miscellany 14: 176. 1996.  A doubtful species.  Basionym: Niebla tuberculata
Vermilacinia tumidula
(Taylor) Spjut ined.
Vermilacinia varicosa Spjut, Sida Botanical Miscellany 14: 176. 1996.
Vermilacinia vesiculosa Spjut, Sida Botanical Miscellany 14: 177. 1996.
Vermilacinia zebrina Spjut, Sida Botanical Miscellany 14: 195. 1996.
Vermilacinia sp. Callao, Peru