Vermilacinia
 

©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
Photos, April 2003
Introduction, Keys and Discussion, February 2005, updated Nov. 2005, revised Sep 2012
image from Follmann (1966) added Nov 2014, Jan 2015 (Images for V. acicularis)
 

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 www.botany.org/bsa/psb/1997/rev25-97.html 

 

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

   Photos of representative specimens for the species in each Subgenus
        Subgenus Vermilacinia
        Subgenus Cylindricaria
   Vermilacinia ceruchis complex in South America
 
 Subgenus Vermilacinia in North America
         Key to the Species of Subgenus Vermilacinia
    Subgenus Cylindricaria
         Key to the Species of Subgenus Cylindricaria
   References
   Nomenclature (authorities, synonyms, and basionyms)

Key to Niebla and Vermilacinia
     

Medulla with string-like (chondroid) strands; cortex with reticulate ridging..................................................... Niebla
Medulla lacking chondroid strands, either with scattered 'knots' of hyphae, or of a single cord; cortex
      mostly a granular sheath or crust.................................................................................................... Vermilacinia

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........................................................ Subgenus Vermilacina

    Cortex membranous, 15-30 μm thick; usually with a regular network
      of areolate depressions; apothecia mostly subterminal and appendiculate...................... Subgenus Cylindricaria

 

Subgenus Vermilacinia

     Subgenus Vermilacinia can be divided into two species complexes that correspond to their disjunct geographical occurrences, a South American V. ceruchis complex with lateral to subterminal apothecia, or without apothecia, and a North American V. combeoides Group with mostly terminal apothecia. In South America, only one species in subgenus Vermilacinia has been recognized (V. ceruchis); however, additional species are evident, some of which may be extinct.  They are further discussed below as variants within a V. ceruchis complex followed by a key proposing their species status.


Vermilacinia ceruchis complex in South America

     Vermilacinia ceruchis is typically terricolous, “lying without adhesion on bare sand” as described by Darwin for a specimen he collected near Iquique, Chile (Peru during his time). His specimen compares favorably with those collected by Joseph Dombey near Lima in Peru and by Archibald Menzies possibly near Valparaiso in Chile  (type collections). This may now be an extinct species (see Follmann 1994).  It is characterized by a thallus consisting of a single (prostrate) cylindrical 'stem' that creeps along the ground with occasional ascending to erect branches, all of which have a relatively thick crusty cortex (75–125 µm).  This is in contrast to a much more divided thallus of thread-like branches as represented by the type for Usnea tumidula from Coquimbo, Chile.  Both variants lack apothecia. 

     The 'tumidula variant' from Chile  (Taylor Herb., FH) is similar to V. ceruchoides from North America that differs by a thallus of more rigid compact branches terminating in short bifurcate branchlets; however, intermediates in South America are evident, although they appear related to the South American species.  Their close similarity was evident to Herre (1906) who stated that he saw the “typical form”  (Ramalina ceruchis) “only once” as “sterile”, occurring sparingly on sandstone cliffs above the sea at Sutro Heights, San Francisco. This was probably what is now regarded as V. ceruchoides  

 

V. ceruchis
Menzies possibly from Valparaiso Chile and Dombey from Lima, Peru
Type, H-Ach 1460
and TLC Data

V. ceruchis, typical.
Iquique Chile
Darwin without number.
Taylor Herbarium (FH)

     Several other variants of the Vermilacinia ceruchis complex in South America differ from their North American counterparts by lateral or subterminal apothecia, while also separated from the related South American species by the thallus branches united at base into a holdfast, presumably as a result of growing on rocks.  Subterminal apothecia are defined as those that develop ± solitarily on upper branches resulting in a change direction of growth from where they originate, or develop an appendage (spur) like branch (especially subgenus Cylindricaria). This in contrast to lateral multiple apothecia on a relatively straight branches as seen in the specimen from Callao.  It is not known whether the Callao variant is extant, although it was reported by Gaudichaud  to be abundant on rocks at Callao (Montagne 1834).  The illustration immediately above is from Montagne (1834, as Usnea ceruchis). Except for the spine-like extension on apothecia, which it seems that Montagne may have added to make it appear more like Usnea, the center and left figures compare closely to the specimen above thought to be from Callao, Peru. The specimen on right that is more branched may have come from spines of cacti or branches of arborescent shrubs at Coquimbo, or possibly terricolous as it resembles V. tigrina from Chile, see image below from Follmann (1966).

     It is interesting to note that specimens in the Tuckerman Herbarium (Farlow Herbarium) shown above—mounted together on one sheet—correspond to Tuckerman's (1882) description and comments for Ramalina combeoides: “quite simple podetiiform thallus, and commonly terminal, now clustered apothecia, grows the the next species in California (Bolander [in reference to R. homalea, Nylander monograph]) but, though certainly marked, is inseparable from South American forms (Tierra del Fuego, Wilkes exp.).”  Thus, Tuckerman (1882) regarded the North American V. combeoides to be the same as saxicolous variants of V. ceruchis, except for the terminal apothecia as he himself indicated.  It should also be kept in mind that V. ceruchis and Niebla homalea were treated in separate genera by Acharius (1810), and that Montagne (1852) combined them into a single genus and species, only to be soon segregated (Trevisan 1861) and later classified into three groups (Bowler 1981, 'homalea', 'combeoides', and tigrina ['ceruchis' misapplied]), and two genera (Spjut 1995).

    

     Subterminal apothecia are seen more often in Subgenus Cylindricaria, usually accompanied by a relatively short continuous branch that ascends sharply upwards as shown above for V. leopardina.  This is in contrast to South American subgenus Vermilacinia with apothecial branches that bend abruptly (geniculate) as evident for the 'roccellaeformis variant'. 

     The mildew-like appearance of the specimens is the result of efflorescence—a chemical change associated with the breakdown of the cortex from which the internal (medullary) hyphae erupt.  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 is apparently rare in lichens, although known in mosses (Physcomitrium).  The mold-like condition develops within a period of six months after a specimen has been collected; 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 (gen. obs., ~9 yrs).   Efflorescence is common in subgenus Vermilacinia  and V. cephalota, V. cerebra, and V. tigrina in subgenus Cylindricaria, in which its occurrence seems closely associated with an unidentified triterpene (T3, Spjut 1996), and with bourgeanic acid, a distinct chemical class of aliphatic depsides (White & Johnson 1994). These compounds—and also the triterpene zeorin—occur in both subgenera but not in Niebla.  Indeed, Niebla and Vermilacinia have little in common in their secondary metabolites except for occasional weak presence of accessory lichen depsidones in Vermilacinia.  The distinction has been further sharpened by finding that species in the Old World 'Bourgeana clade'—also classified in Niebla (Rundel & Bowler 1978; Bowler & Marsh 2004)—are nested within the Old World Ramalina (Sérusiaux et al. 2010).

 

 

Summary: Key to South American species of Vermilacinia subgenus Vermilacinia

1. Basal branches solitary, or without a central attachment point; apothecia lacking; terricolous................................2
1. Basal  branches arising from a common attachment; saxicolous..............................................................................4

     2. Thallus a single cylindrical 'stem', 2–3 mm wide near base, sparingly branched; Valparaiso and
            Iquique Chile, Lima Peru, extinct?...................................................................................................V. ceruchis
     2. Thallus of numerous much-divided branches < 2mm wide..................................................................................3

3.  Basal branches loosely united, wide spreading, long attenuate; Coquimbo Chile, extinct?.....................V. tumidula
3.  Basal branches tightly united, shortly branched, acicular or shortly bifurcate near apex;
          N Am. V. combeoides group........................................................................................................ V. ceruchoides

4.  Basal branches erect, inflated; apothecia lateral; Callao Peru (see also V. vesiculosa, N Am)..........Vermilacinia sp.
4.  Basal branches spreading, not inflated; apothecia subterminal; Coquimbo Chile (see also
           V. subterminalis
), N Am............................................................................................................V. roccellaeformis

 

Subgenus Vermilacinia in North America

     The North American species in the subgenus Vermilacinia are usually recognized by subterminal to terminal, often aggregate apothecia, in contrast to solitary to subterminal or lateral apothecia in the South American species. Included are species without apothecia found on rocks rather than on earth. 

     One species, V. vesiculosa, is recognized for lateral large bladder-like (utricular) swellings and for the pycnidia that develop in tubercular lobes that form the thalline margin.  Bladder-like or gall-like outgrowths ~ 1 mm—that are common on Vermilacinia and also Niebla—contain lichenicolous Heterobasidiomycetes Tremella nieblae (Diederich 2007); however, the development of lateral swellings in V. vesiculosa and the terminal swelling in the corticolous V. cerebra, appear related to an aborted development of apothecia.  The utricular swellings in V. vesiculosa are exceptionally large, ~10 mm diam.

    Four species—Vermilacinia johncassadyi, V. ligulata, V. reptilioderma, and V. rosei—are recognized by a different chemo profile—triterpenes in RF Classes 1–2—than what is typical for the Subgenus Vermilacinia in North America as well as South America.  They occur in southern Baja California Norte, Isla Cedros, Vizcaíno Peninsula and nearby Isla San Roque.  In TLC, these terpenes appear quite strong, and seem to replace terpenes in higher RF classes, but not zeorin and [-]-16 α-hydroxykaurane.   Vermilacinia johncassadyi is morphologically similar to V. laevigata. The type has strongly recurved branches with sessile apothecia along the upper branch margins; V. laevigata, which generally does not occur in Baja California except for one location in the northern chaparral region (~ 15 miles southwest of San Vicente), typically has apothecia on apical short bifurcate lobes.  Vermilacinia ligulata may be compared to V. polymorpha in the Channel Islands in having relatively short basal branches; the Baja California species is ± digitately divided into twisted lobes, whereas the California species has more cylindrical dichotomously divided branches.  Vermilacinia reptilioderma, a relatively rare species, is closely related to V. paleoderma, a common species along the Pacific Coast south of Punta Baja. They are sympatric on the Vizcaíno Peninsula and on Isla Cedros and can only be distinguished with confidence by their lichen substances.  Similarly, V. rosei strongly resembles V. varicosa, in which they are sympatric on Isla San Roque, and V. rosei has also been collected on Isla Cedros.   Except for V. ligulata, these may be regarded as sibling species.

The following key defines the species in the North American Vermilacinia subgenus Vermilacinia.

Key to Species of Subgenus Vermilacinia: North America

Names in plain italics unpublished. Reference to other images made without knowledge of chemical data.

                    

           1     Cortical surface with regular protruding ridges, creases, or crater-like depressions…........................................… 2
           1     Cortical surface smooth, with occasional irregular depressions or ripples or transverse cracks............................. 12

           2     Thallus cushion-like, of short branches tightly compacted, 1–2 (-4) cm high, rarely unbranched for more than
                     10˟ the width of the branch; cortical surface bumpy from protruding knotted hyphae....................................... 3
          
2     Thallus with long spreading branches, usually 2–6 cm long, generally unbranched >10˟ branch width, or
                       with twisted irregularly widened branches; cortical surface appearing creased or glazed.…..............…………. 7

 3(2) Thallus of mostly undivided tubular branches with terminal or subterminal apothecia, the branches arising from 
    a relatively narrow central area; sterile branches sometimes present, obtuse or apically swollen, not abruptly
    narrowed to pointed apex; zeorin often absent; Isla Guadalupe, Santa Catalina Is., peninsular BCN from
    San Quintín to Sonoma Co., CA............................................................................ Vermilacinia combeoides 

 3     Basal branches simple to shortly bifurcate near apex, or densely branched below apex, often matted
together but not united into a common attachment point; branches mostly without apothecia (<10%),  
or entirely absent; spur or isidiod branchlets usually present, reduced in V. pumila.......................
..................... 4

4 (3) Thallus lacking terminal acicular branchlets, or abruptly with narrow acicular isidia; basal branches sparingly
             divided, swollen above base........................................................................................................................5
4      Thallus commonly with terminal acicular branchlets, often shortly bifurcate; basal branches dividing several or
             more times, or intricately divided into threadlike branchlets............................................................................6

5(3)  Basal branches closely compacted, without isidia or soredia, mostly simple and stubby, or shortly divided
           near apex, slightly swollen before tapering to a pointed apex; Isla Guadalupe, Channel Islands,
           peninsular BCN north of Ensenada to Alameda Co., CA.................................................... Vermilacinia pumila
5 (4) Basal branches spreading, irregularly divided, isidia or soredia or both present, especially towards blunt apex,
           isidioid branchlets often present; Channel Islands; Santa Monica Mts...............................Vermilacinia acicularis


6      Thallus branches <1 mm wide; Channel Islands and peninsular BCN from near San Vicente to Marin Co.,
          CA........................................................................................................................ Vermilacinia ceruchoides   
6.     Thallus branches 1–5 mm wide; Morro Bay, CA...........................................................Vermilacinia tuberculata

7(2) Branches with bladder-like swellings; pycnidia elevated on pustular lobes; rare, between
                    Punta Canoas and Puerto Catarina, peninsular  BCN ....................................................Vermilacinia vesiculosa
         
7     Branches without bladder-like swellings; pycnidia flush with cortical surface...............................……………….... 8

          8(7) Primary branches 5–10˟ longer than wide, twisted, lobulate; triterpenes present in RF classes 1–2; Isla
                   Cedros and adjacent peninsular BCN................................................................................ Vermilacinia ligulata
         
8     Branches ±linear (>10˟ longer than wide ), cylindrical-prismatic (in x-section), not lobulate; triterpenes
                   present or absent in RF Class 1–2.................................................................................................................... 9

          9(8). Branches flexuous, 0.5–1.5 mm wide; cortex on upper branches thinner, eroded and pitted;
                        Isla Cedros and adjacent peninsulas.....................................................................… Vermilacinia cedrosensis
         
9.     Branches curved, straight or geniculate, 1–5 mm wide; cortex not eroded on upper branches, the surface
                        plicate to honeycomb-like........................................................................................................................  10

10(9).  Apothecia aggregate on expanded marginal and terminal lobes constricted below; endemic to 
                NW Isla Cedros……......................................……………………….....…….....…..... Vermilacinia convoluta
10       Apothecia solitary to aggregate on terminal cylindrical to expanded lobes (generally more flattened), 
                 not notably constricted................………………………………………......................................................
11

11  Triterpenes present in RF Classes 1–2; Vizcaíno Peninsula and Isla Cedros…... 
        ................................................................................................................................. Vermilacinia reptilioderma
11  Triterpenes absent in RF Classes 1–2; widespread, Baja California,
         Channel Islands..………………………………………..............................................….. Vermilacinia paleoderma

12(1)  Branches distinctly blade-like, two-edged......................................................................................................... 13
         
12      Branches subcylindric to flattened and dilated ................................................................................................. 15

          13(12) Triterpenes present  in RF classes G: 1–2; rare, Isla Cedros, southern half of
                        peninsular BCN, also evident on Santa Rosa Is (Sharnoff photo) and at  Pt Lobos, CA
                        (Parrish photo)....................................................................................................  Vermilacinia johncassadyi
          13       Triterpenes absent in RF classes: 1–2............................................................................................................. 14

          14(13)  Branches uniformly blackened around base to slightly above, usually less than 4 cm long, 1–4 mm
                        wide;
apothecia cupular; rare, Northern Vizcaíno Desert ................................................... Vermilacinia rigida

          14(13) Branches irregularly blacked near base and above, usually >3.5 cm long and 4–7 mm wide;
                         apothecia lenticular; Channel Islands, northern peninsular BCN to
                          northern CA.............................................................................................................. Vermilacinia laevigata

          15(13) Branches uniformly cylindrical, irregularly blackened above base, often with black transverse bands or cracks........ 16
         
15       Branches dilated or inflated, blackened mostly near base, or not at all blackened, not transversely cracked............. 17

16(15) Apothecia mostly terminal; Californian coastal chaparral region, San Francisco to San Quintín,
                       Channel Islands............................................................................................................... Vermilacinia procera
          16      Apothecia subterminal, much like corticolous V. leopardina; rare, rocks near the ocean,
                        Bahía de San  Quintín............................................................................................ Vermilacinia subterminalis

17(15) Branches dilated, flattened and/or digitately divided; Baja California.................................................................... 18
         
17       Branches inflated or deflated, mostly simple; California and Baja California......................................................... 19

18(17) Triterpenes present in RF Classes 1–2; Isla San Roque and Isla Cedros...................……..……...  Vermilacinia rosei
         
18       Triterpenes absent in RF Classes 1–2; Isla San Roque..........................................………….. Vermilacinia varicosa

19(17) Branches inflated, ± round in x-section; Isla Guadalupe, Channel Islands, peninsular BCN from
                        Punta Banda to southern CA............................................................................................. Vermilacinia robusta
          19       Branches deflated, with longitudinal folds or intermarginal ridges; infrequent, Channel Islands,
                        rare in BCN (Punta Cono)........................................................................................... Vermilacinia polymorpha

 

V. acicularis, San Clemente Island
Between Eel Point and Seal Cove,
Santesson 19746, 16 Apr 1966, Holotype (US)

                                                                                    

 

acicularis-10147.jpg (26340 bytes)

V. acicularis
San Clemente Island
Bratt 10147, Oct 1997

V. acicularis
Los Angeles  Co.
Santa Monica Mountains, Conejo Mountain, Jason Hollinger 196947
19 Oct 2011
From Wikipedia, where identified Niebla isidiaescens

cedrosensis-10923.jpg (53906 bytes)

V. cedrosensis
E of Vizcaíno Peninsula, San Francisco Mts (BCS), Spjut & Marin 10923, Apr 1989

cedrosensis-10549.jpg (76557 bytes)

V. cedrosensis
Isla Cedros,
Spjut & Marin 10549A, Apr 1989

 

V. cedrosensis
San Andrés Ranch, BCN
Spjut & Marin 9069, May 1985

V. cedrosensis
Vizcaíno Peninsula,
NW of Bahía Tortugas (BCS)
Spjut 9689a, May 1986

V. ceruchis
Chile
Darwin s.n.
Taylor Herbarium (FH)
 

V. ceruchis variant
Chile, Hassler Expedition (FH)
(Ramalina roccellaeformis)


 

V. ceruchis variant
 Callao Peru? (FH)

 

ceruchoides-9045.jpg (52985 bytes)

V. ceruchoides
Cerro Solo, BCN
Spjut & Marin 9045, Apr 1985

combeoides-10030.jpg (62194 bytes)

V. combeoides
Left:
Marin Co.. Pt. Reyes, CA
Spjut 10030, May 1987
Right: Probably California, mounted on sheet with other specimens from Wilkes Expedition to
Tierra del Fuego (FH)

V. combeoides
Pt. Lobos State Park,
image, Chris Parrish 151683
no chemical data

V. combeoides
Pt. Lobos State Park,
image, Chris Parrish 151687
no chemical data

San Francisco Bay Area,
4 images, Stephen Sharnoff

V. convoluta
NW Isla Cedros
Spjut & Marin 10555, Apr 1989

 

 

V. johncassadyi
Isla Cedros, BCN
topotype, Apr 1989

 

 

johncassadyi--10532.jpg (24065 bytes)johncassadyi-11531.jpg (29073 bytes)

V. johncassadyi
Is. Cedros (left) Punta Cono (right)
BCN; Spjut & Marin 11531, Apr 1990

 

V. laevigata
Pt. Lobos St Park,
Monterrey Co., CA
No chemistry data
image, Chris Parrish 69358

Santa Rosa Is, San Francisco area,
oo., CA
No chemical data. N laevigata1, 2,5
images, Stephen Sharnoff

laevigata-9047C.jpg (39036 bytes)

V. laevigata
Cerro Solo, BCN
Spjut & Marin 9047C, Apr 1985

ligulata-10542.jpg (37860 bytes)

V. ligulata
Isla Cedros
Spjut & Marin 10542, Apr 1989

ligulata-9074L.jpg (64800 bytes)

V. ligulata
San Andrés Ranch, ~100 km N of Guerrero Negro, Spjut & Marin 9074L, topotype, May 1985

ligulata-11434.jpg (54084 bytes)

V. ligulata
between Punta
Canoas and Punta Blanca, BCN
Spjut & Marin 11434, Apr 1990

paleoderma-12668.jpg (58347 bytes)

V. paleoderma
near Punta Baja, BCN
Spjut & Marin 12668, Mar 1993

V. paleoderma
Isla Cedros, Cabo de San Angustín,  Nash 34611, Mar 1994

V. paleoderma
Punta Cono, BCN
Spjut & Marin 11528, Apr 1990

V. paleoderma
near Punta Rocosa, BCN
Spjut 10321, Mar 1988

V. paleoderma
near Punta Rocosa, BCN
Spjut 10296, Mar 1988

V. paleoderma
near Punta Rocosa, BCN
Spjut 10317, Mar 1988

V. paleoderma
near Punta Rocosa, BCN
Spjut 10325, Mar 1988

V. polymorpha
Punta Cono, BCN
Spjut & Marin 11526B

Spjut, R. 2000. Notes on lichen Vermilacinia polymorpha (Ramalinaceae) and related species in Baja California, Mexico. IV. Symposium on botany research in Baja California and adjacent areas areas. Ensenada, B.C. Mexico, Sep. 13–17.  Image extracted from poster presentation.

 

procera-11932.jpg (88438 bytes)

V. procera
Bahía de San Quintín, BCN
Spjut & Marin 11932, Feb 1991

 

V. pumila
Pt. Lobos State Park
Monterrey Co., CA
Chris Parrish, 2011-5-19 (151682)

pumila-9027A.jpg (17148 bytes)

V. pumila
La Misión, between Tijuana
 and Ensenada, BCN
Spjut & Marin 9027, Apr 1985

 

reptioloderma-isotype.jpg (89258 bytes)

V. reptilioderma
Punta  Eugenia, BCS
 Spjut 9734, isotype, May 1986

 

rigida-9975.jpg (64970 bytes)

V. rigida
El Marrón ridge
S of Punta Negra, BCN
Spjut & Marin 9975,
isotype, May 1985

 

robusta-9035.jpg (44747 bytes)

V. robusta
Punta Banda, BCN
Spjut & Marin 9034, Apr 1985

V. rosei
Isla Cedros, precipitous rocks, northwest coast
Spjut & Marin 10535, Apr 1989
 

 

V. vesiculosa
Punta  Camachos, BCN
holotype 

 

Subgenus Cylindricaria

 

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

cerebra-9722.jpg (103565 bytes)

V. cerebra
Near Rosarito, BCN
Spjut 9722, 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

cerebra-9796.jpg (128477 bytes)

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

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)

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

 

vermilacinia_howei_9722.jpg (84534 bytes)

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

 

V. leonis
Chile, Santiago
Looser 838 (FH)

 

leonis-9606D.jpg (117478 bytes)

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

 

leonis-12692.jpg (185568 bytes)

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

leopardina9334.jpg (82766 bytes)

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

leopardina-9986-isotype.jpg (66912 bytes)

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

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

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

 

 

 Subgenus Cylindricaria

     Vermilacinia Subgenus Cylindricaria is easily recognized in North America by the thallus divided into cylindrical branches from a holdfast along with its habit of growing on bark of shrubs or trees.  The thallus branches have a relatively thin membranous cortex (15–30 µm) covering a loose network of medullary hyphal cells that are longitudinally oriented, converging at various intervals into knotted bundles. This network arrangement allows for the collapsing and expansion of the cortex onto the knotted hyphae as related to desiccation (dissipation) and hydration from fog.  Subgenus Cylindricaria is further characterized by having black pycnidia, or in some species colorless (sterile) pycnidia worm-like branches, absence of medullary chondroid strands, presence of black pycnidia or black spots/bands, and subterminal appendiculate apothecia.

     Two common species, V. leopardina and V. corrugata, are recognized by their differences in the folding of the cortex as evidently related to zonal moisture differences correlated with the production of the diterpene [-]-16 α-hydroxykaurane.  For example, in the Northern Vizcaíno Desert of Baja California, Vermilacinia leopardina, which has the diterpene, is mainly seen along the immediate coast, in contrast to the inland V. corrugata that lacks the diterpene.  The cortical surface of the former species generally appears smooth, whereas in the latter, the cortex is strongly rugose.  However, on the Vizcaíno Peninsula (Southern Vizcaíno Desert), the character features in this relationship become reversed.  The smooth cortical features are seen in the more coastal Vermilacinia howei, which lacks the diterpene, in contrast to the rugose thallus of V. nylanderi that contains the diterpene.

    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 in contrast to lateral soralia seen in the other species.

     The South American species include chemotypes not found in North America. Vermilacinia flaccescens, which was treated broadly to include thalli with a honeycomb-like cortex and others with a firm smooth cortex (Spjut 1996), was distinguished by the presence of methyl 3,5 dichlorolecanorate, a depside that has also been reported in species of Ramalina.  This secondary metabolite may have been acquired from hybridization with South American species of Ramalina.  Similar thalli having this depside but lacking the terpenes generally seen in North American Cylindricaria are considered Vermilacinia cactacearum (Follmann) Follmann & Werner (Schedae ad Lichenes Exsiccati).

     Another South American ramalinoid species, Niebla nashii, which was described to have terpenoid chemistry (Sipman 2011), except perhaps not the diterpene ('no bloom'), also has methyl 3,5 dichlorolecanorate; however, it does not appear to belong to Vermilacinia or Niebla.  It lacks pycnidia as well as apothecia (Sipman 2011).  While it was also described to have medullary chondroid strands free from the cortex, and its overall resemblance to Ramalina lacera (With.) J. R. Laundon was noted, the presence of more than one depside suggests affinities to the Bourgeana clade of Ramalina (Krog, H. & H. Østhagen. 1980; Sérusiaux et al. 2010).  However, it may also be noted that Vermilacinia lacera (With.) Follm. & Wern. has been proposed (G. Follmann & B. C. Werner: Sched. Lich. Exs. Univ. Coloniensis Ed., Fasc. 24, 11. 2003. No. 479).

     The evolutionary ties of North American subgenus Cylindricaria to South American species are evident in the sorediate species (V. leonis) and the non-sorediate V. cerebra and V. leopardina as already indicated under subgenus Vermilacinia.  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 in South America includes 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 evolutionary trend in Cylindricaria seems to be loss of secondary metabolites, which may have occurred after Cylindricaria were established in both North and South America.  The reversed pattern in the diterpene chemistry between the Vizcaíno Peninsula and northern peninsula species also indicates that their features evolved at a time when the Vizcaíno Peninsula was perhaps an island separated from the main Baja peninsula. 

 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 (Niebla) 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.
4          
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
8
        Cortical surface deeply folded and/or regularly lacunose; black
transverse bands
                  or enlarged spots absent or not regularly present; mostly Channel
Islands
                  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

 

References

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.

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.

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
cedrosensis
, is described from Baja California, Mexico. Phytologia
76: 458-460.

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
6:497-499.

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.

_________. 1996. Niebla and Vermilacinia (Ramalinaceae) from California and Baja California. Sida, Botanical Miscellany 14: 1–207, 11 plates.  

_________. 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.

Trevisan, V. G. 1861. Ueber Atestia eine neue Gattung der Ramalineen aus Mittel-Amerika. Flora 4:49-53.

Tuckerman, E. 1882. Synopsis of the North American Lichens Pt 1. S.E. Cassino, Publishers, Boston.

White, J. D. and A. T. Johnson. 1994. Synthesis of the aliphatic depsides +(-) bourgeanic acid. J. Org. Chem. 59: 3347–33558

 

Nomenclature

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 ined.
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 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