Vermilacinia
 

©The World Botanical Associates Web Page
Prepared by Richard W. Spjut
Photos, April 2003
Introduction, Keys and Discussion, February 2005, updated Nov. 2005
 

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 Subgenera of Vermilacinia
   Photos of representative specimens for the species in each Subgenus
        Subgenus Vermilacinia
        Subgenus Cylindricaria
   Discussion on the Taxonomy of Vermilacinia
     
 Species Concept in Subgenus Vermilacinia
         Key to the Species of Subgenus Vermilacinia
       Species Concept in 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; 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

                                                                                    

acicularis-10147.jpg (26340 bytes)

V. acicularis
San Clemente Island
Bratt 10147, Oct 1997

cedrosensis-10549.jpg (76557 bytes)

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

cedrosensis-10923.jpg (53906 bytes)

V. cedrosensis
E of Vizcaíno Peninsula, San Francisco Mts (BCS), Spjut & Marin 10923, 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
Lima, Peru
Type, H-Ach 1460
TLC Data
 

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

V. ceruchis
Chile, Hassler Expedition (FH)

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

 

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: From Argentina
Tierra del Fuego
Wilkes Expedition (FH)

johncassadyi--10532.jpg (24065 bytes)

V. johncassadyi
Isla Cedros, BCN
Spjut & Marin 10532,
topotype, Apr 1989

johncassadyi-11531.jpg (29073 bytes)

V. johncassadyi
Punta Cono, BCN
Spjut & Marin 11531, Apr 1990

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

procera-11932.jpg (88438 bytes)

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

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

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.

 

Discussion on the Taxonomy of Vermilacinia
 

     A key to subgenus Vermilacinia was first drafted in late 1986 as part of a revision of the Niebla complex.  At that time, Spjut had already recognized most species of Vermilacinia, including those later published by Marsh and Nash (1994), Bowler et al. (1994) and Riefner et al. (1995), which were no doubt inspired by Spjut's manuscript already in circulation since 1990. 

     Although Spjut pursued his study of Niebla beginning in 1986 with the support of the Smithsonian Institution and Dr. Hale, Phillip Rundel objected in reply to an inquiry letter by Spjut, because Dr. Rundel indicated he was still working on Niebla. Spjut relayed this information to Dr. Hale who indicated that Dr. Rundel was not likely to have the time to do anything further in the genus, and neither was Peter Bowler since he was teaching in Canada.  Spjut, therefore, decided to pursue Niebla.

    Thomas Nash III at ASU was made aware of Spjut's Baja California collections and his study of Niebla through communications from Mason Hale and from Spjut who met Dr. Nash at the Smithsonian during 1987–88.  Dr. Nash subsequently initiated a Greater Sonoran Lichen Desert Project.

    A paper submitted by Spjut for peer review in 1990 was criticized by reviewers for not citing material from more herbaria; Spjut's collections from the Baja California mainland were undoubtedly the largest available at the time, while he had also cited the US collections, small loans obtained by Mason Hale from herbaria at Harvard and types from European herbaria.  Although Dr. Hale felt that Spjut did not need additional loans (possibly because he did not want Spjut's taxonomy to be influenced by annotations at other herbaria), Spjut subsequently obtained loans from Santa Barbara (Charis Bratt) and Colorado (Weber at COLO), but had to commute from Laurel MD to Washington DC to study the loans, while the material he had collected was readily available except for the entire WBA herbarium of subgenus Vermilacinia (1985–1986 collections) that was to remain at the US at the request of Dr. Hale who had also studied the specimens in regard to the 1986 draft key mentioned above.

      Unfortunately, Dr. Hale developed cancer and following his death in 1990, Nash and others began to jump in on the study of Niebla.  Spjut's manuscript was already in press when the quickie taxonomy papers produced by Nash's associates began to appear in Phytologia and Mycotaxon.  The loan from Santa Barbara had already been returned, while new annotations to specimens from US and COLO were provided.  Because the presentation of species names in the paper was partly in alphabetical order with references to figures, a lot of small changes had to be made.  And it was felt necessary to review the types at ASU even though some species were obviously the same as those that had been described by Spjut.  Had Spjut's paper been allowed to proceed to completion without interjection of others, there would have been less taxonomic confusion today in the genus.  It might be noted that Spjut was a member of the American Bryological and Lichenological Society and the British Lichenological Society.  Supposedly, societies work towards a common goal such as to promote a systematic research on lichen taxonomy.  But from Spjut's experience it would appear that lichenologists tend to make taxonomy and its research more difficult for its members as well as for the students.  It certainly appears as one of the least studied fields of organisms.

     Most confusion that has resulted from the sudden appearance of papers on Niebla taxonomy is confined to Subgenus Vermilacinia.  Ironically, the species in this subgenus are the easiest to comprehend and no doubt this is why Bowler, Nash and their associates have recognized more North American species in this subgenus (8 species) than in Niebla (3 species) or subgenus Cylindricaria (2 species).  Another reason for more species in this subgenus is that one of the two centers of species diversity recognized by Spjut (1995, 1996) lies relatively close to the taxonomists in southern California as evidenced by the type locations for Vermilacinia ceruchoides from Ventura Co., V. laevigata from San Luis Obispo Co.,  V. tuberculata and V. procera from Morro Bay, and V. polymorpha from Santa Catalina Island, except for Marsh and Nash recognition of V. cedrosensis from Isla Cedros. 

     Nevertheless, Spjut had independently reached the same conclusions as Rundel and Bowler for their new species of V. procera (= Vermilacinia robustiella Spjut ined.), except Spjut disagrees with the inferred circumscription based on geographical distribution given that would appear to include V. paleoderma.  Charis Bratt (and probably others) apparently had also recognized this as possibly a distinct species as evident from a specimen she had sent Mason Hale in 1985 or 1986 inquiring as to whether it was Niebla homalea.

     The basionym for Vermilacinia ceruchoides, was based on an unpublished name (nomen nudum) that had appeared in a number of publications for some time (Bowler et al. 1994), and from specimens on loan, Spjut adopted this name and designated an appropriate type specimen, one collected by Santesson from San Clemente Island (Spjut ms 1990–1994), whereas Bowler et al. (1994) chose a type that seemed less relevant to the origin of the name.  Spjut had also recognized another similar species, V. pulvinata (Spjut ined.) based on a type he designated from Cerro Solo in BCN.  As a result of the type designated by Rundel & Bowler (Bowler et al. 1994) for V. ceruchoides, the Santesson specimen no longer fit Spjut's concept of V. ceruchoides, and as a result it became the type for the other species, which Spjut decided should get another name, V. acicularis, distinguished by the isidia or isidia-like branchlets.  Vermilacinia tuberculata was described by Riefner et al. (1995, as Niebla tuberculata) to have similar features, which are more conspicuous in its type; however, they reported that this species was deficient in the diterpene [-]-16 α-hydroxykaurane, and possibly lacking in zeorin as well.  The absence of the diterpene [-]-16 α-hydroxykaurane in species of Vermilacinia had only been known to Spjut (1996) to occur in Cylindricaria, not in subgenus Vermilacinia.  Therefore, Spjut (1996) considered V. tuberculata a distinct species.

      However, upon a review of another specimen collected by Riefner at Morro Bay, Riefner 86-30 at COLO, annotated V. tuberculata by Spjut on loan to US, it had been determined to contain the diterpene with trace amounts of the triterpene zeorin.  This specimen was cited by Spjut (1996) under V. ceruchoides.  Thalli deficient in zeorin also had been noted by Spjut (1996) for V. combeoides and V. rigida.  The “isidiose-papillate” (or “tuberculate”) “thallus” was reported by Riefner et al. (1995) as the key distinguishing feature of this species.  A type at ASU was informally recognized by Spjut and Marsh (Spjut visit to ASU, Apr 1996) to differ mainly by size, generally larger than V. ceruchoides and V. acicularis.  Riefner et al. (1995) indicated that V. tuberculata was abundant around Morro Bay Rock Reserve, but it was not indicated whether V. ceruchoides also occurred there.

     Bowler and Rundel is the authority for the basionym of V. laevigata (Bowler et al. 1994), a species that Spjut had earlier recognized as V. halei (Spjut ined., 1986–1995), whereas Rundel is the first author for two other species they described (N. ceruchoides, N. procera).   Unlike V. ceruchoides and V. procera, there was no indication from specimen loans or literature as to when this species was recognized by Bowler.  Bowler (1981), in a anatomical review of the Ramalinaceae, indicated in his summary that “most species” in the  Niebla homaela group have “conspicuous chondroid stands in the medulla which are not attached to the cortex.”  He did not say what the exceptions were, while Spjut (1996) does not know of any exceptions. Spjut had conducted his own independent anatomical studies of many hundreds of specimens of the Niebla complex, the results of which were illustrated on packets that were submitted to the US herbarium, later replaced by clean packets, and detailed in descriptions (Spjut draft ms, 1990–94).  Based on anatomical examination of more than 1,000 specimens, the chondroid strands are clearly a key character for distinguishing Niebla from Vermilacinia. 

     Bowler  et al. (1994) also had carved out other species that still need study.  One is V. polymorpha they recognized to occur in the Channel Islands (Santa Catalina Is.) and nearby mainland (Orange and Ventura Cos. ), and another is V. cedrosensis from Isla Cedros and nearby Vizcaíno Peninsula.  The intervening region of 500 miles of coastline was thought to have only V. laevigata and V. procera (Bowler et al. 1994), but these were Californian species according to Spjut (1996).  Spjut (ms 1990–96) had considered V. polymorpha and V. cedrosensis to be variants of V. paleoderma, and that more study was needed, although Spjut's (1986 draft key) had earlier segregated Vermilacinia (Niebla) paleoderma into four species that Dr. Hale questioned.  Among the questionable species was V. albicans (Spjut ined.), that became formally established as V. cedrosensis Marsh & Nash (1994).  Support for maintaining this species was evident to Spjut from 25 g samples collected for cancer research of V. cedrosensis and V. paleoderma from the San Andrés Ranch in May 1985 (Spjut & Marin, 9069, 9074), treated as distinct species in the field, and other specimens collected for floristic studies in 1986 from the Vizcaíno Peninsula.  The flexuous branchlets that characterize this species appear related to slight differences in cortical ridges that form the support framework; in V. paleoderma, the main ridges frequently divide diagonally, whereas in V. cedrosensis the ridges run lengthwise with more frequent traverse ridges.

     As for V. polymorpha, Bowler et al. (1994) indicated this to be an inland Californian species, in contrast to V. robusta and V. paleoderma that were recognized by Spjut to occur along the immediate coast.  Also, Spjut (1996) recognized Vermilacinia paleoderma to occur mostly in Baja California south of San Quintín; however, collections from the Channel Islands were cited by Spjut (1996) as belonging to this species.  There is also confusion between V. paleoderma and V. polymorpha; for example, Nash 36411, shown above, was identified by Nash as Niebla polymorpha; however, in Bowler et al. (1994) and Bowler and Marsh (2004) this keys to V procera.   Part of the confusion is due to Bowler et al. (1994) failure to show the type and provide an adequate description. In Bowler’s description of V. polymorpha, he did not indicate the thallus size or length of the thallus branches, only a width to about 6 mm.  The cortical features mentioned by Bowler et al. (1994), “surface crinkled and irregularly lacunose” along with their “representative specimen” (instead of the type) indicate that their concept of this species is limited to that under the V. robusta lineage described below.   Additionally, Bowler & Marsh (2004) treated V. paleoderma and V. ligulata as synonyms of V. laevigata, a species recognized for having a relatively smooth cortex in contrast to the Nash specimen shown above.  Rather than producing a quickie taxonomy paper for Phytologia, Spjut (1996) provided a comprehensive review of the Niebla complex that required more time for peer-review.  The misapplication of names by Bowler & Marsh 2004), the misidentifications of Niebla photos in their publications (Bowler & Marsh 2004), and other inconsistencies in the taxonomic treatment as a whole (Bowler & Marsh 2004), are unconscionable; Bowler et al. (1994) and Bowler and Marsh (2004) should be regarded “Opera Utique Oppressa” for inclusion in the International Botanical Code of Nomenclature.

     Based on Californian species, Spjut's taxonomy was more conservative than that of Bowler et al. (1994), and who knows just how many species they might have recognized if Spjut's (1996) paper had not appeared.  Despite the confusion in subgenus Vermilacinia, the lumping by Bower & Marsh (2004) of many of Spjut's (1996) species of Niebla and of subgenus Cylindricaria  (Spjut 1995, 1996) under the names found in Bowler et al. (1994), and their treatment of Niebla pulchribarbara as a synonym of N. josecuervoi (Bowler & Marsh 2004), makes it easier for the novice to work with the two drastically different, alternative taxonomic treatments.

Species Concepts and Keys to the Species


Subgenus Vermilacinia

     The following key to the species of Subgenus Vermilacinia reflects an understanding of a more natural arrangement compared to a more practical approach published in Sida Miscellany (Spjut 1996).  The species are seen as belonging to four different lineages based on cortical differences, summarized as follows.

      1. A combeoides lineage with five or more species in California and South America characterized by having a fine closely recticulate cortex with bulging knots of hyphae along cortical ridges.  Vermilacinia ceruchis, misapplied  by Bowler et al. (1994) and Bowler & Marsh (2004) to corticolous species in North America, is typically a terricolous species of coastal deserts in South America. The type from Peru has branches that lie flat on sand, an adaptation that might be compared to the cactus Stenocereus eruca, an endemic to the Pacific coast of southern Baja California, recognized as a distinct species for its prostrate stems.  Other forms of V. ceruchis are similar to the North American Vermilacinia ceruchoides, while a saxicolous form resembles V. combeoides, which also occurs in South America.  Vermilacinia ceruchis is distinguished by the relatively long branchlets or ‘spur shoots,’ in contrast to lack of such branchlets in Vermilacinia combeoides.   Other species, V. acicularis and V. ceruchoides, have needle-like spurs.  In V. pumila the spur branchlets are further reduced, although this may have been acquired from hybridization between V. ceruchoides and V. combeoides

          2. A Vermilacinia robusta lineage is recognized by expanded lobes with broad elliptical to round depressions between cortical ridges.  This lineage includes 4–5 species occurring mainly on the islands off the Pacific Coast of southern California and Baja California.   Vermilacinia robusta is identified by its inflated branches, in contrast to two other island species that have flattened branches, V. rosei and V. varicosa.  A species found more on the mainland, Vermilacinia procera, has regularly shaped branches as in V. combeoides, but its cortical and medulla features are similar to those of V. robusta.  One other questionable species, Vermilacinia polymorpha, might be interpreted as a hybrid complex of species involving V. robusta as the main thalloid form.  This includes forms that appear transitional to V. paleoderma (Spjut 1996, 1997), a species with a more regularly plicate cortex, and forms that have a blade-like morphology of V. laevigata and V. ligulata.  Its branches may be characterized as tubular to blade-like, obovoid, usually from 3–6 (or more) mm in diam., with a contorted or wrinkled surface.  Based on the type and representative specimen (Bowler et al. 1994), and excluding forms with reticulate cortical ridging (Spjut 1997), V. polymorpha may be recognized as a deflated morph of V. robusta. 

      3. A third group is characterized by a regularly plicate cortex in which the surface appears glazed with folds along reticulate ridges and also depressed between the ridges and often cracking along a pleat.  The ancestral species is believe to have features similar to Vermilacinia ligulata, broad reticulate ridging, twisted branches, and triterpenes in RF Classes 1–2.  Vermilacinia ligulata occurs along the east coast of Isla Cedros and the nearby peninsula of Baja California Norte, but not on the Vizcaíno Peninsula.  Species such as V. paleoderma and V. reptilioderma may have evolved from hybridization between an ancestral V. ligulata and V. robusta; for example, V. paleoderma may have acquired the typical “ceruchoid” chemistry, and V. reptilioderma the linear branches. 

      4. Finally, a fourth group is recognized by blade-like branches with a glazed cortical surface, typified by V. laevigata, a species that was long overlooked and included under Niebla homalea.  This confusion still persists in Bowler et al. (1994) and Bowler & Marsh (2004) treatments because they on one hand argue that Niebla cannot be distinguished from Vermilacinia by the presence of chrondroid strands, while they do so anyway in their keys to species.  No specimens or species were cited to support their contention that the kind of chrondroid strands seen in Niebla are found in Vermilacinia.  Other species of the laevigata group are V. rigida and V. johncassadyi.  These differ by the blackened area around the base of the thallus; V. rigida has relatively narrow short branches, while V. johncassadyi has longer oblong to obovate branches and the triterpene chemistry found in the ligulata group mentioned above.

Key to Species of Subgenus Vermilacinia

           1     Cortical surface closely reticulately ridged or lacunose …..……………..............… 2
           1     Cortex surface smooth to broadly depressed, ridges with rounded angles.............. 12

           2     Areolate surface defined by very close, narrow, fine ridges, with transverse
                    cracks, the ridges uneven, protruding and bumpy (from knotted hyphae).............. 3
          
2     Cortical surface appearing plicate or glazed...........………....…..............…………. 7

 3(2) Thallus of mostly simple branches without spurs, arising from a holdfast;
 apothecia terminal; zeorin often absent; Isla Guadalupe, Santa Catalina Is.,
 peninsular BCN from San Quintín to Sonoma Co., CA; also in South
 America, Tierra del Fuego, Argentina........................... Vermilacinia combeoides

3    Spur branchlets usually present, reduced in V. pumila; thallus often with
                    numerous branches densely compacted in low rounded
mats, usually
                    without a holdfast and < 2 cm high (except S America)...................................... 4

4(3) Branches and spurs usually long; apothecia oriented perpendicular to branches
           (lateral); thallus loosely attached to substrate in terricolous forms, with
           spreading branches from a holdfast in
saxicolous forms; coastal Peru and
           Chile.............................................................................
Vermilacinia ceruchis

4.   Branches all short; apothecia terminally aborted or absent; thallus of densely
 compacted branches in low rounded mats......................................................... 5

5(4). Branches mostly simple, 0.5-2 mm wide, abruptly and bluntly  pointed at
          apex, obtuse, rarely shortly bifurcate near apex;  Isla Guadalupe, Channel
          Islands, peninsular BCN north of Ensenada to southern CA... Vermilacinia pumila

5    Thallus of dichotomously divided, or intricately divided, capillary or threadlike
                    branchlets, 0.7 mm or less wide, shortly bifurcate to acuminate apex..................  6

6(5) Thallus isidiate, or with tiny isidialike branchlets, and sorediate;
          Channel Islands ...........................................................  Vermilacinia acicularis

6.   Isidia or soredia lacking, branchlets all spur-like; Channel Islands and peninsular
                    BCN from near San Vicente to Marin Co.,CA.............. Vermilacinia ceruchoides

7(2) Branches with bladder-like swellings from aborted development of apothecia; rare,
                   between Punta Canoas and Puerto Catarina,
                   peninsular BCN..............................................................Vermilacinia vesiculosa
         
7     Branches without bladder-like swellings..............………………………………….... 8

          8(5) Primary branches oblong, strap-shaped with twisted branchlets; triterpenes present
                   in RF class 1–2; Isla Cedros and adjacent peninsular BCN...... Vermilacinia ligulata
         
8(3) Branches linear and cylindrical prismatic (in x section); triterpenes
                   present or absent in RF Class 1–2....................................................................... 9

9(8). Branches flexuous, 0.5–1.5 mm wide; main cortical ridges mostly longitudinally
         oriented with transverse connections and rounded angles, cortex often whitish

         and pitted; Isla Cedros and adjacent peninsulas..............… Vermilacinia cedrosensis
9      Branches geniculate, 1–5 mm wide; cortical ridges diagonal and transverse,
         with sharp acute angles, or if rounded then plicate, the surface plicate to
         alveolate.........................................................................................................  10

10.  Cortical ridges sharply raised from surface, prominently alveolate; endemic to 
         NW Isla Cedros……...…………………………….....…..... Vermilacinia convoluta
10  Cortical ridges glazed and/or plicate................………………………………………. 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) Branches 1–4 mm wide, closely fastigiate; Northern Vizcaíno Desert...............
                 
.............................................................................................. Vermilacinia rigida
         
13       Branches 3–7 mm wide, widely spreading apart................................................. 14

          14(13) Branches irregularly blackened from base to apex, mostly 3–6 cm long;
                         triterpenes absent in RF classes: 1–2; apothecia lenticular; Channel Islands,
                         northern peninsular BCN to northern CA........................ Vermilacinia laevigata

14      Branches uniformly blackened around base to slightly above, usually less than
                        4 cm long; apothecia cupular; triterpenes present  in RF classes G: 1–2;
                        Isla Cedros, southern half of peninsular BCN...........  Vermilacinia johncassadyi

          15(13) Branches uniformly cylindrical, irregularly blacked from base to apex................... 16
         
15      Branches dilated or inflated, blackened mostly near base, or not at all blackened.... 17

16(15) Apothecia terminal; primarily a Californian species, from San Francisco to
                        San Quintín, Channel Islands............................................ Vermilacinia procera
          16       Apothecia subterminal; rare, rocks near the ocean, Bahía de San  Quintín....
                  ................................................................................... Vermilacinia subterminalis

17(15) Branches dilated, flattened and digitately divided towards apex; 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 and Isla Cedros........
                        ..................……………………………………………….. Vermilacinia varicosa

19(17) Branches inflated; Isla Guadalupe, Channel Islands, peninsular BCN from
                        Punta Banda to southern CA............................................. Vermilacinia robusta

19       Branches strongly wrinkled, or deflated; infrequent in Channel Islands,
                        rare in BCN (Punta Cono).......................................... Vermilacinia polymorpha

         

 Subgenus Cylindricaria

      Speciation in the Subgenus Cylindricaria reflects adaptation to ecological differences in moisture.  Key characters include the presence or absence of the diterpene [-]-16 α-hydroxykaurane and the depside, methyl 3,5 dichlorolecanorate.

     In the Northern Vizcaíno Desert of Baja California, the occurrence of the diterpene correlates with proximity to coastal influence of fog and thallus morphology.  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 cortex of the former species is generally smooth, whereas the latter is strongly rugose.  However, on the Vizcaíno Peninsula (Southern Vizcaíno Desert), the chemical features are reversed in regard to thallus proximity to the coast.  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.

     Methyl 3,5 dichlorolecanorate, a relatively uncommon depside in lichens, was recognized only in    Vermilacinia flaccescens, a South American species.  This secondary metabolite was perhaps acquired from hybridization with South American species of Ramalina.  Thalli having this depside but lacking the terpenes usually found in Cylindricaria are considered Vermilacinia cactacearum.

     Other key features are the presence of depsidones and bourgeanic acid.  The evolutionary trend in Cylindricaria seems to be loss of secondary metabolites, which appears to have occurred after Cylindricaria species were established in both North and South American.  The reversed pattern in the diterpene chemistry between the Vizcaíno Peninsula and northern peninsula species also indicate that their features evolved at a time when the Vizcaíno Peninsula was perhaps an island separated from the main Baja peninsula.

     Another taxonomic feature of Cylindricaria is the development of soralia.  Four or more species are recognized to have this feature, in contrast to one species recognized by  Bowler & Marsh (2004).  It is interesting  that the sorediate species, and two other nonsorediate species, V. cerebra, and V. tigrina, produce bourgeanic acid and depsidones.  The morphology of V. cerebra appears intermediate to subgenus Vermilacinia.

Key to Species of Subgenus Cylindricaria 

1(0).     Thallus with methyl 3,5 dichlorolecanorate; South America.
               
1a With
[-]-16 α-hydroxykaurane........................................ Vermilacinia flaccescens
               
1a Without
[-]-16 α-hydroxykaurane................................. Vermilacinia cactacearum
1.         Thallus lacking methyl 3,5 dichlorolecanorate; North and
South America.......................  2

2(1).    Thallus with distinct orbicular 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.......
                 ......................................................................................... Vermilacinia cephalota
3.         Branches all of uniform width, < 1.0 mm wide; acicular near apex; cortex pale in
                 color, yellowish-green or straw-colored.................................................................. 4

4(3).     Thallus regularly dichotomously divided, shortly bifurcate near apex, with or without
                  black bands and spots
                     4a Thallus with well defined black spots or transverse bands; California, especially
                          common in the Channel Islands...................................... 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; Baja California
                  Sur and 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 immediatecoast......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.

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.

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.

 

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 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: 166. 1996.
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 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 varicosa Spjut, Sida Botanical Miscellany 14: 176. 1996.
Vermilacinia vesciculosa Spjut, Sida Botanical Miscellany 14: 177. 1996.
Vermilacinia zebrina Spjut, Sida Botanical Miscellany 14: 195. 1996.