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 Sér 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 Sér 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.
