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mNK Technical Report NMFS Circular 446

Marine Flora and Fauna of
the Northeastern United States.
Lichens (Ascomycetes) of
the Intertidal Region

Ronald M. Taylor

August 1982

QK

586.4

.T39

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atoiospheric Administration

National Marine Fisheries Service

NOAA TECHNICAL REPORTS

National Marine Fisheries Service, Circulars

The major responsibiHlies of the National Marine FIshenes Service (NMFS) are to monitor and assess the abundance and geographic distnbution of fishery
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tion subsidies. It collects, analyzes, and publishes statistics on vanous phases of the industry

The NOAA Technical Report NMFS Circular series continues a series that has been in existence since 1941 . The Circulars are technical publications of
general interest intended to aid conservation and management. Publications that review in considerable detail and at a high technical level certain broad areas of
research appear in this series. Technical papers originating m economics studies and from management investigations appear in the Circular series.

NOAA Technical Report NMFS Circulars are available free in limited numbers to governmental agencies, both Federal and State They are also available in
exchange for other scientific and technical publications in the marine sciences Individual copies may be obtained (unless otherwise noted) from D822, User
Services Branch, Environmental Science Information Center, NOAA, Rockville, MD 20852 Recent Circulars are;

418. Annotated bibliography of four .Atlantic scombnds; Scomberomonn hranl-
iensis. S. cavalla, S. maculaius. and S. rcgatis. by Charles S. Manooch III, Eugene
L Nakamura, and Ann Bowman Hall. December 1978, iii + 166p.

419. Marine llora and fauna of the northeastern United States Protozoa Sarco-
dina: Amoebae, by Eugene C. Bovee and Thomas K. Sawyer January 1979, in +56
p., 77 figs. For sale by the Superintendent of Documents, U.S. Government Printing
Office, Washington, D C 20402, Stock No OO.VO 17-00433-.^

420. Preliminary keys to otoliths of some adult fishes of the Gulf of Alaska,
BeringSea, and Beaufort Sea, by James E. Morrow. February 1979, in +32 p., 9 pi.

421 Larval development ol shallow water barnacles of the Carolinas (Cimpedia:
Thoracica) with keys to naupliar stages, by William H Lang Februarv 1979,
iv+39p., 36 figs , 17 tables

422 A revision of the catsharks, family Scyliorhinidac, by Stewart Springer April
1979, vtI42 p.. 97 figs. For sale by the Superintendent of Documents, U.S. Gov-
ernment Printing Office, Washington, DC 20402, Stock No. 003-020-00147-5.

423 Marine flora and fauna ol the northeastern United Slates Crustacea Cuma-
cea, by Les Watling. April 1979. iii+23 p.. 35 figs. For sale by the Superintendent
of Documents. US Government Printing Office. Washington. DC. 20402, Stock
No 003-017-00446-5.

424. Guide to the leptocephali (Elopiformes. AnguiUilormes. and Notacanthi-
formes). by David G. Smith July 1979. iv ^39 p.. 54 figs

425 ,Marinc flora and launa ol the northeastern United States. Arthropoda: Cim-
pedia, by Victor A. Zullo. April 1979, in +29 p., 40 figs For sale by the Superin-
tendent of Documents, U.S. Government Printing Office. Washington. DC. 20402.
Stock No 003 017-00453-8.

426. Synopsis of biological data on the rock crab. Cuncer irninini\ Say. b\
Thomas E. Bigford. May 1979. v +26 p.. 1 1 figs. 21 tables

427. Ocean variability in the V S Fisherv Conservation Zone. 1976. by Julien R
Goulcl. Jr. and Eli/aheth D Havnes. editors July 1979. iii + ^62 p

428. Morphological comparisims of North American sea bass larvae (Pisces: Ser-
ranidael. by Arthur W, Kendall. Jr August I979,iv+50p . 43 figs. 9 tables

429. Synopsis of biological data on tunas of the genus Eullnmui\. by Howard O.
Yoshida. October 1979, iv ^57 p., 40 figs , 30 tables.

430. Guide to identification of some sculpin (Cottidae) larvae from marine and
brackish waters off Oregon and adjacent areas in the northeast Pacific, by Sally L.
Richardson and Betsy B. Washington. January 1980, iv +56 p.. 32 figs,. 45 tables,

431. Guide to some trawl-caught manne fishes from Maine to Cape Hatteras,
North Carolina, by Donald D. Flescher March 1980. iii+34 p,

432 Synopsis of biological data on bonitos of the genus Sarda. by Howard O.
Yoshida May 1980, iv+50p.,43 figs.. 26 tables.

433. Synopsis of biological data on stnped bass. Monme vavofifa (Walbaum). by
EileenM. Setzler. WalterR Boynton. Kalhryn V. Wood. Henry H. Zion. Lawrence
Lubbers. Nancy K. Mountford. Phyllis Frere. Luther Tucker, and Joseph A.
Mihursky June 1980. v+69p.. 9 figs.. 24 tables.

434. Osteology, phylogeny. and higher classification of the fishes of the Order
Plectognathi (Tetraodontiformes). by James C. Tyler October 1980. xi+422 p .
326 figs.. 3 tables.

435 Field guide to fishes commonly taken in longline operations in the western
North Atlantic Ocean, by Joseph L Russo. January 1981. iii+51 p .58 figs.

436 Synopsis of biological data on frigate tuna. Aims ihazard. and bullet tuna. A.
rochei. by Richard N. Uchida. January 1981. iv+63p., 52 figs., 27 tables.

437 Fishery publication index. 1975-79. by Lee C Thorson May 1981. iii + l 17
P

438. Manne flora and fauna of the northeastern United Stales. Scleractinia. by
Stephen D. Cairns. July 1981. in + 15 p.. 16 figs.. 2 tables

439 Marine llora and launa of the northeastern United States. Protozoa: Sarco-
dina: benthic Foramimfera. by Ruth Todd and Dons Low. June 1981. iii-r5l p. 23
figs.

440 Marine flora and fauna of the northeastern United States. Turbellana: Acoela
and Nemertodermatida. bv Louise F Bush July 1981.iii+71 p . 184figs

44 1 , Synopsis of the biology ot the swordfish. Xiphms gladuis Linnaeus, by B, J.
Palko, G- L. Beardsley, and W. J. Richards, November 1981. iv+21 p.. 12 figs,. 5
tables.

442. Proceedings of the Sixth U.S.— Japan Meeting on Aquaculture. Santa Bar-
bara. California, August 27-28. 1977. by Carl J Sindermann (editor). March 1982.
iii +66 p

7 /\

./

NOAA Technical Report NMFS Circular 446

^OWMOS^,.

Marine Flora and Fauna of
the Northeastern United States.
Lichens (Ascomycetes) of
the Intertidal Region

Ronald M. Taylor

August 1982

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U.S. DEPARTMENT OF COMMERCE

Malcolm Baldridge, Secretary

National Oceanic and Atmospheric Administration

John V, Byrne, Administrator

National Marine Fisheries Service

William G. Gordon, Assistant Administrator for Fisheries

FOREWORD

This NMFS Circular is part of the subsenes "Manne Flora and Fauna of the Northeastern United
Stales." which consists of original, illustrated, modern manuals on the identification, classification, and
general biology of the estuanne and coastal manne plants and animals of the northeastern United Slates.
The manuals are published at irregular intervals on as many taxaof the region as there are specialists avail-
able to collaborate in their preparation

Geographic coverage of the "Manne Flora and Fauna of the Northeastern United States" is planned to
include organisms from the headwaters of estuanes seaward to approximately the 200 m depth on the con-
tinental shelf from Maine to Virginia, but may vary somewhat with each major taxon and the interests of
collaborators. Whenever possible representative specimens dealt with in the manuals are deposited in the
reference collections of major museums of the region.

The "Manne Flora and Fauna of the Northeastern United States" is being prepared in collaboration
with systematic specialists in the United States and abroad. Each manual is based pnmanly on recent and
ongoing revisionary systematic research and a fresh examinatin of the plants and animals. Each major
ta.xon. treated in a separate manual, includes an introduction, illustrated glossary, uniform originally illus-
trated keys, annotated checklist with information when available on distribution, habitat, life history, and
related biology, references to the major literature of the group, and a systematic index.

These manuals are intended for use by biology students, biologists, biological oceanographers.
informed laymen, and others wishing to identify coastal organisms for this region. Often they can serve as
guides to additional information about species or groups.

The manuals are an outgrowth of the widely used "Keys to Manne Invertebrates of the Woods Hole
Region." edited by R. I. Smith in 1964. and produced under the auspices of the Systematics Ecology Pro-
gram. Marine Biological Laboratory. Woods Hole. Mass. After a sufficient number of manuals of related
taxonomic groups have been published, the manuals will be revised, grouped, and issued as special vol-
umes, which will consist of compilations tor phyla or groups of phyla.

CONTENTS

Introduction 1

Methods of collection and studying 1

Ecology 2

Glossal^ 4

lUusti^ated key to species 6

Annotated list of species 15

Selected bibliography 22

Systematic index 24

Acknowledgments 25

Coordinating Editors comments 26

The National Marine Fisheries Service (NMFS) does not approve, rec-
ommend or endorse any proprietary product or proprietary material
mentioned in this publication. No reference shall be made to NMFS, or
to this publication furnished by NMFS, in any advertising or sales pro-
motion which would indicate or imply that NMFS approves, recommends
or endorses any proprietary product or proprietary material mentioned
herein, or which has as its purpose an intent to cause directly or indirectly
the advertised product to be used or purchased because of this NMFS
publication.

Marine Flora and Fauna of the Northeastern

United States. Lichens (Ascomycetes) of

the Intertidal Region

RONALD M.TAYLOR'

ABSTRACT

Thi> manual ircals (he lichens lound in ihc intertidal region trom New Jersey to Newloundland Methods of
collection, preparation, and study are briefly treated. Twenty-two species are covered, both in an illustrated key
and an alphabetical listing, witfi brief descriptions and notes on ecology and dislribution. A glossary of terms is
included

INTRODUCTION

METHODS OF COLLECTION AND STUDYING

The content and style of this paper are intended to encourage
and serve all who might wish to study the lichens of the fascinating
and complex marine littoral environment. It is evident from the lit-
erature and from inquines directed to the author that the manne
lichen flora presents an interesting enigma to biologists of diverse
disciplines. It is hoped that this paper will help them in studying,
recognizing, and accurately reporting the lichens of the littoral
zone.

A lichen is not "a plant" but rather a life <ir growth form com-
prised of two kinds of plants, a fungus and an alga, growing
together in a symbiotic relationship. Although botanists differ as to
the exact nature of the symbiotic relationship, each member of the
relationship seems to benefit from the other In reality, there are
probably different kinds of relationships in different lichens. Most
lichens are fungus dominated, i.e.. the shape and most of the mass
of the lichen is derived from the fungus. All the manne lichens
treated in this manual, except for the genus Lichiiui. are fungus
dominated and the fungus is an Ascomycete.

Manne lichens occur in three growth forms: fruticose. foliose.
and cru.stose . Fruticose forms are usually upnght or pendant in rela-
tionship to the substrate; they stand out away from the substrate.
Lichiiia. the only genus of fruticose lichens in the littoral region
grows in a shrublike form. Foliose forms, as the name implies,
have a leafy appearance, often in rosette form, and may be removed
from the substrate. The only genus of foliose lichens in the littoral
region is Xanthoria. Crustose forms often appear as discolored
areas on rocks and are usually destroyed if one attempts to remove
them from the substrate. They may be shiny or dull, black, green,
or brown. Most marine lichens are crustose and grow on rocks: one
species, Anhopyrenia halodyles. grows on barnacles.

Lichens may be found in a continuous range of habitats, from
low tide to mountain tops. The species included in this manual are
those found in the intertidal zone plus the normal spray zone but not
what is often called the "storm spray zone."

Some of the lichens in this manual are obligate marine lichens,
i.e., they grow only where they are struck directly by saltwater
Others are facultative, being found also in areas remote from the
sea. My criterion for including a facultatn c lichen is its occurrence
with, or at the same height on the shore as, an obligate marine
species.

'Science Depariment. Lansing Conimuniiv College. Lansing. MI 48914

Manne lichens inhabit rocks or the barnacles attached to them.
Most littoral lichens are crustose and cannot be removed from the
rocks without being destroyed. They must be collected along with a
piece of the rock substrate, using rock chisels and a 3-lb hammer
Ideally one should strive to collect specimens on pieces of rock
small enough to fit well into 4" x 6" herbarium packets. This will
not always be possible because the lichen may grow on an irregular
piece of rock that does not break easily. Further preparation can be
done in the laboratory, but it is usually easier to reduce the size of a
rock in the field.

Once the specimen is removed it must be placed in a suitable
container. Paper bags are quite satisfactory if the bag is strong.
Unfonunately. the specimens are often wet, and the wet bag may
need to be placed in another bag. Plastic or other waterproof bags
are not desirable since they retard drying and promote molding. If a
waterproof bag is used in the field, the specimen should be transfer-
red to a paper bag and air dried within a day of collection.

A rock saw would seem to provide a fairly ideal solution to the
problem of shaping rocks for herbarium packets in the laboratory.
However, the use of a rock saw requires a liquid coolant to protect
the blade. In preparing lichen specimens, water should be used
rather than the oil employed in lapidary work. This works well with
nonlittoral specimens, but with littoral lichens the soaking that
results from the water coolant and the additional water required to
rinse the specimen cau.ses excessive reticulation that is detrimental
to futjire study.

Hydraulic devices are available which break rocks by applying
pressure to chisellike jaws. These produce fractures in the rocks
that are not as neat or predictable as the cut of a saw but they do
avoid reticulation and are quite satisfactory.

After sizing, the rocks should be glued to cards of suitable size.
This prevents abrasion of the specimens and degradation of the
rock. Also, it provides a way to permanently identify the rock frag-
ments examined for the lichen in question.

Many of the diagnostic characteristics of littoral lichens are
readily observed with a dissecting microscope at a magnification of
20 power Bnght illumination is important because of the dark color
of many thalli, especially in the genus Verrucana. Thalli should be
observed in both wet and dry conditions. Water added to the thallus
sparingly to avoid unnatural reticulation often increases its trans-
parency, thus revealing subsurface patterns.

Sections of fruiting structures may be made by hand with a
razorblade. The small size of the perithecia of many species of Ver-
ntcaria are difficult but not impossible to section. A double edged
razorblade. broken lengthwise, is most suitable.

Microtome sections are complicated by the thin crustose nature
of the thalli and their adherence to the rock substrate. Furthermore
the involucrella of pyrenocarpous lichens are hard and brittle and
do not respond well to the usual paraffin imbedding techniques.

Two techniques employing a freezing microtome are satisfac-
tory. One technique involves pouring a shallow layer of agar in a
small container The perithecium is moistened several times with
water containing a small amount of wetting agent, such as alcohol.
The perithecium and the subtending and surrounding thallus are
then scooped from the rock with a razor blade, taking care to lift the
entire penthecium from the rock. Faulty diagnosis will result if the
base of the perthecium is left on the rock. The thallus and perithe-
cium are then transferred to the surface of the agar A drop of melted
agar placed over the lichen material embeds it between the tw o lay-
ers of agar A cube cut from the agar is then placed on the freezing
microtome so that the thallus is perpendicular to the blade and the
vertical axis of the penthecium is parallel to the blade. The matenal
is then frozen and sectioned.

The other technique involves commercially available synthetic
plastic matenals intended for embedding frozen sections. A mound
is built up on the microtome's object disc and frozen. A vertical face
is cut facing and perpendicular to the blade. The lichen material to
be cut is prepared as before and then pressed firmly against the
embedding matenal. The pressure melts the plastic and when the
pressure is released the plastic and embedding lichen fuse and
freeze solid.

Both techniques have the disadvantage of leaving visible resi-
dues on the slide surrounding the sections. This is a problem only if
the slide is to be photographed.

Some species of lichens give a color reaction to potassium
hydroxide. A drop of 10% solution is applied with a dropping
pipette or capillary tube, directly to the cortex, or to medulla
exposed by fragmentation or an incision.

Santesson (1939) described a number of environmentally
induced morphological vanations in littoral lichens in general and
described in detail the variations of certain selected species. Both
Weber (1962) and Santesson (1939) have emphasized the impor-
tance of understanding the range of vanation induced by the
environment.

The author has traced variations of single thalli from continu-
ous to areolate and from smooth to ridged. The author has also
observed variations of penthecia from sunken to prominently
raised on a single thallus. Even the darkness of the lower side of the
excipulum (inner lining of perithecium) is quite variable within a
single species. Shade modification is well known and predictable
among littoral lichens. Shade causes lighter pigmentation and
reduction in size and number of juga (black pegs, lumps, or ndges
protruding from the surface). Some species that are distinct in
appearance when growing in good illumination appear much less
distinct when growing in the shade. The distinguishing features of a
species are frequently reduced by the shade modification. As a
result there is considerable taxonomic confusion among the shade
modifications of Verrucaha striatiila, V. dilmarsica, and, to a
lesser degree. V. erichsenii. Too much stress can be placed on
perithecium size. Variations of perithecium size on a single thallus
beyond the range accepted in some taxonomic keys have been
observed. Thus the taxonomic character "perithecia slightly
smaller (or larger) than . . " should be regarded with suspicion.

If one's study is restricted to a few specimens, it is easy to see
dichotomy among the species examined. As the number of speci-
mens increases, the kinds and degrees of variations also increase
and extreme forms are bridged by lesser variations. Since the litto-
ral environment is one of great physical contrasts and many micro-
habitats. it is not suprising that considerable taxonomically
confusing variations in the lichens have resulted.

ECOLOGY

Between low and high tide marks the rocks of the shore are
largely covered by macroaglae such as Fiicits sp. Among the hold-
fasts of such algae, the manne lichens compete for space with bar-
nacles. Bryozoa. and such algae as Hihienhrantia and
Lithothamnia . The lichen Anhopyrenia luilodyles often occupies
the shells of the barnacles with which the other lichens compete.
Penthecia. the cone-like fmiting stmctures of marine lichens, are
often seen protruding through the thalli of Hildenbrantia or Lilho-
ihamnia which have overgrown them. Such competition with the
lichen Vernicana mucosa has mixed results; either the lichen or the
alga may establish itself on top. Only the lichen genera Anhopyre-
nia. Sligmidium . and Vernicaria are found throughout this zone of
the shore. The Vernicaria species inaura and ceiilhocarpa do not
extend very far down into this zone.

From the high tide mark upward for about 0.5 m is the barnacle
belt. The barnacles are so successful in the competition for space
that only their shells are available as a substrate for lichens. The
lichen Anhopyrenia haUxlytes is found here in pits dissolved into
the barnacle shells.

Extending upward from the barnacle belt is the black belt pop-
ulated by various species of Vernicaria in competition with blue-
green algae, especially of the genus Calothrix.

Investigators in northern Europe find a well defined orange
belt above the black belt and a white "RamaUna belt" above that.
In northeastern North America the orange belt is intermittent or
absent and the "Ramalina belt" is entirely absent. The orange belt,
where present, is compnsed of members of the genera Caloplaca
and Xanthoria. Lichina confinis. a black fmticose lichen, may be
found in this belt or extending down to the black belt. Various other
lichens may extend into the orange belt but in the few and incom-
plete orange belts encountered in northeastern North America by
the author, only Lecanora granlii. a gray cmstose lichen w ith cup-
like fruiting bodies, was found. Nothing above the orange belt was
considered in this study because it is the highest zone to contain
obligate marine lichens.

The lichen flora collected at the northern end of the study
range was comprised of 22 species. At the southern end of the range
the lichen tlora was reduced to two species. In an attempt to explain
the distribution in terms of environmental cause, a multiple regres-
sion analysis was used to assess the impact of environmental factors
on the success of mdividual species and on the general diversity of
species. The environmental factors whose individual and interac-
tive effects were computed are: salinity, air and water temperatures,
solar insolation, and tidal range. No simple answers emerged either
for the success of any species or for general diversity. However, by
tallying the number of times that each environmental factor was
involved as one of the most important influences, the following fre-
quencies resulted: salinity, 21; insolation, 15; air temperature, 12;
tidal range, 7; and water temperature, 2.

Where sufficient collections were made to provide adequate
data, vertical distnbutions of the various marine lichens were
graphed (Figs. 1-8). In these graphs the extent above mean high

-, 10m r-

50":
H

Figure 1.— Vertical distribution of Arlhupyrenia halodytes.

Figure 4. — \ertical di*>tribution of Verrucaria erichsenii.

<-

ahrs
enrs

12hrs

20hr5
24hrs

Figure 2. — Vertical divlribulion of \'erriuaria degelii.

Figure 5. — Vertical distribution of Verrucaria maura.

-I 10m ,-

Figure 3. — \ertital distribution of Verrucaria ditmarsica.

Figure 6. — Vertical distribution of Verrucaria micrmpora.

ihr

Rhi-

;.„,. ^

Figure 7. — N'erlical distribution of Verrucaria mucosa.

Figure 8. — \ertical distribution of Verrucaria striatula.

tide is given in meters. Due to the great vanation in the distance
between mean low tide and mean high tide (ca. 1 to 10 m). the dis-
tance below high tide is given in hours of submergence. Each inter-
val on the X-axis represents 10% of the total collections of that
species by the author In each figure the graph on the right repre-
sents the vertical distnbution on exposed shores and the graph on
the left represents the vertical distribution on sheltered shores.

GLOSSARY

The use of lichenological terminology in this publication has
been reduced to a minimum. Several terms are illustrated as well as
defined.

Apical At the tip or apex (Fig. 9).

Apiillifciiiiii (pi. Apoihc'ciii) Disc, saucer, or cup shaped faiit-
ing body (Fig. 9).

Areolaie Broken by clefts into small, usually angular and irreg-
ular patches (areoles).

AivdIc (pi. Areoles) Area of a thallus divided from the rest of
the thallus by fissures (Fig. 10).

Arete (pi. Aretes) Sharp peak elevated above a ridge.

Ascocarp (pi. Ascoccirps) An Ascomycete fruiting body giving
nse to and containing asci and ascospores (Figs. 9. 10).

Asiiis (pi. -4.VC7) A sack-like stmcture within which ascospores
are formed and contained in the ascocarp (Fig. 10).

Carhniuueoiis Made up largely of carbon deposits.

lamina! soredia

-phycobiont
medul la

thall ine
ma rg i n

upper cortex

lower cortex

paraphyses-

hymenium

Figure 9.— Composite foliose lichen vwth apotlu'cia.

Citrine Lemon yellow color.

Conidium (pi. Conidia) Asexual reproductive .spores of Asco-
mycete.

Continuous Thallus unbroken by gaps or fissures.

Cortex (pi. Cortices or Corlexes) The compact outermost fun-
gal layer of a lichen thallus (Figs. 9. 10).

Craterous Having a depression resembling the crater of a vol-
cano.

Crenulale Having small rounded lobes.

Crustose A type of thallus which usually adheres tightly to the
substrate and cannot be removed from it without destruction of the
thallus: lacks a lower cortex and rhizines (Fig. 10).

Cusf> (pi. Cusps) Crescent shaped mound.

Dendritic Having a pattern resembling the branching of a tree
limb.

Dispersed Scattered.

Endolithic Growing within the rock substrate.

Epilhecium (pi. Epithecui) Covenng of the hymenium.

Evufiinate To protrude as if a pocket turned inside-out.

Excipulum (pi. Excipida) As used here, the inner wall of a
peritheciuni.

Foliose A type of thallus, leatTike or lobed. often with rhizines
or hairlike processes on lower surface, usually with lower cortex,
algal layer limited to upper side (Fig. 9).

Frulicose A type of thallus that is shrublike, stalked, or pen-
dant with the algal layer usually on all sides.

Fusiform Narrowed both ways from a swollen middle.

Glabrous Having a smooth even surface.

Globose Spherical .

Holdfast (pi. Holdfasts) The basal portion of an algal thallus
attaching it to the substrate.

Hyaline Colorless.

Hymenium (pi. Hymeniu) The layer of the ascocarp containing
the asci and usually paraphyses (Fig. 9).

Hypothallus (pi. Hypothalli) A marginal outgrowth of hyphae
from the thallus of a crustose lichen (Fig. 10).

Immersed Sunken mostly or entirely below the surface of the
thallus.

Impressed Indented

Involucrellum (pi. Involucrella) The usually black and carbo-

naceous covering of a peritheciuni (Fig 10)

Isidium (pi. Isidia) Small cylindncal, coralloid, or globose
outgrowths of the thallus serving as vegetative dis,seminules. They
are covered with cortex and contain both fungal and algal compo-
nents (Fig. 9).

Isthmus (pi. Isthmuses) A narrow passage.

Jugum (pi. Juga) Protrusions of carbonaceous material extend-
ing at least partly above the thallus surface as ridges, pegs, or irreg-
ular lumps (Fig. 10).

Luminal On the flat surface of the thallus (usually upper) (Fig.
9).

Lenticular An area, pointed at each end, inclosed by two inter-
secting convex curves.

Littoral lichen Growing between the tides. The term is
extended herein to include those struck by saltwater due to wave
action and is u.sed interchangeably with the term "manne."

Locule (pi. Locules) Chamber or cavity.

Medulla (pi. Medullae) An area of loosely packed hyphae
internal to the thallus and its projections such as podetia, isidia, etc.
(Figs. 9, 10).

Morpholype (pi. Morphotypes) A thallus of a given species
having a different appearance than other members of the species.

Necral Occurring after death.

Necral reticulation A network of cracks caused by treatment
and/or storage after collection.

Nodular convex Like a small rounded lump.

Oblong ellipsoid In the form of an elongated circle; more elon-
gated than the typical elipse.

Oblong ovoid In the form of an elongated oval.

Olivaceous Having a color similar to that of a green olive.

Orbicular Spherical or nearly so.

Ostiole (pi, Ostioles) The apical opening of a perithecium (Fig.
10).

Paraphysis (pi. Paraphyses) Sterile hyphae intermingled with
asci in the hymenium of an ascocarp (Figs. 9, 10),

Parasymbiont (pi. Parasymbiunts) An organism living in
harmless association with another without mutual benefit.
Pendant Dangling.
Perithecoid In the shape of a perithecium.

perithecium

hypotha

areol

ucrell um

pegs
jugum

ortex

paraphyses

•-medul la

Figure 10, — C'ontpnsiU' crustose lichen vvidi periltiecia.

Perithccium (pi. Periihccia) A t'lask-shaped ascocarp enclosed
except for the apical pore (ostiole). It is usually more or less sunken
into the thallus (Fig. 10).

PhycobioM (pi. Phycohioms) The algal member of the lichen
symbiosis (Figs. 9, 10).

Pseudosepium (pi. Pseudoscpta) Appearing to be an mternal
wall but not composed of the same matenal forming the cell wall.

Pycnidium (pi. Pycnidia) Small tlask-shaped to globular struc-
tures resembling penthecia but containing conidia instead of asci
and ascospores.

Pyrencocarpous Having penthecia.

PyrenoUchen (pi. Pyrenolichens) Lichen having penthecia
(Fig. 10).

Reniform Kidney shaped.

Reticulation A netlike appearance.

Rimose Having minute cracks, usually referring to a form of
areolation.

Saxicolous Growing on a rock or rock-like substrate.

Sessile Not supported by a stalk.

Simple Describing a spore having a single, undivided cavity.

Siirediare Having soredia on the thallus.

Sorediiini (pi. Soredia) A means of vegetative reproduction of
the lichen consisting of masses of hyphae and their associated algae
rupturing through and not covered by or containing corte.x (Fig. 9).

Spray zone (pi. zones) That area above high tide that is struck
by drops of seawater resulting from the impact of waves on the
rocky shore under nonstorm conditions.

Storm sprav zone (pi. zones) That area, above the usual spray
zone, that is sprayed with seawater only during a storm.

Subconvex Barely curved outward.

Subejfuse Having boundanes that are pooriy distinguished.

Submerged Sunken entirely or mostly below the upper surface
of the thallus.

Submoniliform Having constrictions at intervals giving an
appearance ncariy like a stnng of beads but not so pronounced.

Tluilline margin (pi. margins) Apothecial nm having the same
composition as the thallus (Fig. 9).

Tliallus (pi. Tlmlli) The lichen body containing both algal and
fungal components.

Tiiberciilale Having a warty appearance.

ILLUSTRATED KEY TO SPECIES

The key is based primarily on charactenstics that can be observed with a dissecting microscope at 20 x magnification; in some cases
observation of spores with a compound microscope may be required.

The vertical sections of perithecia of Verrucaria species illustrated in the key are typical sections.

Because of the extreme degree of variation of species of manne lichens (especially of Verrucaria). identification of species from the key
alone should be regarded as tentative until the more detailed descnption in the Annotated List of Species is read.

Five species from the Canadian Maritime Provinces are included even though at present they are not known from the eastern United
States. At least one species. Verrucaria maura. has not been collected in the eastern United States recently but the author has seen specimens
collected from the area over 50 yr ago.

1 Thallus foliose (Fig. 9); always orange and KOH + purple 2

1 Thallus crustose (Fig. 10) only in a few species orange and KOH + purple 4

1 Thallus erect, fruticose, composed pnmanly of algal (Caloihnx) filaments; ascocarp penthecioid (Fig. II) Lichina conjinis

Figure \\.— Lichina confinis. Scale is O.I
mm.

2 (/) Thallus with .soredia mostly apical (Fig. 12l. thallus lobes finely divided l0,2-0,5 mm wide) .

.Xanilioria cundclnria

Figure 12.— Thallus iif Xantliuria candclaria. .Scale is
I nini.

/r^^-^^i^^WwW

^-

2 (1) Thallus not sorediate, lobes w idcr than 0.? mm 3

3 (2) Lobes Hattcncd and 1 mm or more broad (Fig. 13) Xanthona panctina

Figure I.V— Thnlliis of Xanthoria parivtina. Si'uk' is 1
rum.

3 {2) Lobes nodular-convex, lobes less than 1 mm wide (Fig. 14) Xanthona elef^ans

Figure 14.— Thallus of Xanthoria elegans. Scale is
mm.

1^

is-~r

4 (/) Ascocarp an apothecium (Fig. ^). thallus light (shades of white, gray, or yellow orange) 5

4 (/) Aseoearp a pentheeium (Fig. 10); occasionally more or less immersed with only ostiole showing; thallus dark

(shades of green, brown, or black) or apparently absent 9

5 (4) Thallus whitish to gray, KOH + yellow; spores colorless, one-celled (Fig. 13), 11-18 v 6-8. 5^m Lecanoni gninlii

Figure 15.— One-celled spiire. .Stale is
lOiim.

5 {4] Thallus yellow orange. KOH + puiple; spores colorless, polarilocular (Fig. 16) 6

Figure 16.

ularihieuhir sport. .Stale is
lO/im.

6 (5) Thallus covered with globose isidia (Fig. 17); spores 9-14 x 4-6;im Culophica vemiculifei

Figurt 17.— Thallus of Caloplaca verruculifera.
Scale is lU mm.

6 (5) Thallus without isidia 7

7 (<S) Thallus forming circular patches with radiating lobes (Fig. 18); spores 10-15 x 5-7;(m Caloplaca scopularis

Figure 18.— Thallus of Caloplaca scnpularis
Scale is 10 mm.

7 (6) Thallus irregular, composed of scattered lobes or numerous granules .

8 {7) Thallus citrine of granules 1-3 mm diameter (Fig. 19). spores 10-18 x 5-8f(m Caloplaca inicrothalUna

F'igure 19.— Thallus of Caloplaca microthalUna.
Scale is 10 mm.

1 (7) Thallus orange yellow, marginal lobules subconvex oriented in radiating manner; .spores 10-14 x 5-7;im (Fig. 20)

Caloplaca marina

Figure 20.— Thallus of Caloplaca marina. Scale is
I mm.

I I

9 (4) Spores one-celled 11

9 (4) Spore^ more than one-celled 10

10 (9) Growing on ihallus of Vernicahiw: paraphyses gelatinizing, spores 10-15 x .^.S-S/jni with 4 locules and distinct cen-

\n\ septum (Fig. 21 ) Stigmidium marinum

Figure 21. — Spore of Sligrrudium
marinum. .Scale is lO/im.

/

\

1
J

\

10 (9) On barnacles or rock; thallus scant to evidently lacking; paraphyses persistent, spores 9-20 x 4-7^in with two locules

(Fig. 11) Anhopyrenia halodytes

Figure 22.— Spore of Anhopyrenia halodytes.
Scale is lO/im.

1 1 (9) Thallus with pegs and/or ridges present (Fig. 23) 16

Figure 23.— a. Thallus with pegs. b. Thallus «ilh ridges. Scale is 1 mm.

a I ♦ b

1 1 (9) Thallus smooth; pegs and ridges lacking 12

12 (//) Thallus areolate (di\ided mto fields by fissures. Fig. 24 1 15

Figure 24. — Areolale thallus. Scale is
1 mm.

I (

12 (//) Thallus continuous or discontinuous but not areolate (Fig. 25) 13

-> .'

c?

Figure 25.— a. Continuous thallus. b. Discontinuous thallus. Scale is
I mm.

13 (/2) Perithecia elevated, often shiny, dome-shaped to hemispherical or sometimes pointed (Fig. 26) 14

Figure 26.— Thallus with ele\ated perithecia.
Scale is I mm.

13 (12) Penthecia submerged to slightly elevated (Fig. 27); spores 8-1 1

4-5 ^m; thallus thick, green to greenish-black
Vcniiccirla mucosa

Figure 27. — Thallus with submerged to
slightly ele\ated perithecia. Scale is I mm.

14 ( /.^) Spores 6-1 1 x 3-5;im .
14 (7i) Spores 18-22 -x 8-9/mi.

. Venucaria micrcyspora
. . . Vernicana silicictla

15 {12) Areoles light brown on dark brown prothallus (Fig. 28); penthecia brown; spores thick walled (11.5) 15-23

(27) X (4) 6-8( 14)/im Vernicana inicniign-scens

Figure 28.— a. Vertical section of
perilhccium. Scale is 0.1 mm. h.
Thallus of Xi'micaria inlernigres-
ceiis. Scale is 1 mm.

15 {12) Prothallus not evident; thallus prommently fissured; penthecia brown or black; spores thin-walled. 9-12 x 5-6f(ni

(Fig. 29) Venucaria ceutlmcarpa

I H

ro^i

Figure 2*). — a. Wrtiial section
of perithelium. Stale is O.l
mm. h. Thallus of Verrucaria
ceuthocarpa. Scale is 0. 1 mm.

10

16 (//) Thallus areolatc (thallus divided into fields by fissures. Fig. 30) 17

Figure 30. — Areolate thallus. Scale is I mm.

16(//) Thallus not areolate 20

17 (76) Areoles bordered by raised juga (Fig. 31).

.18

Figure 31 . — Areoli-s bordered b)
juga. Scale is 0.1 mm.

17 iI6) Areoles not bordered by juga (Fig. 32) .

.19

Figure 32. — Areoles wilhout juga.
Scale is 0. 1 mm.

11

18 {17) Perithecia usually immersed, ostiole border frequently raised, spores 9-12 x 5-6/im; juga rarely limited to borders of

areoles, usually scattered within areoles (Fig. 33); thallus more translucent wet than dry Verrucaria degelii

Figure 33.— a. Vertical section of
perilheciuni. Scale is 0.1 mm. b.
Thallus of Verrucaria degelii.
Scale is 1 mm.

18 (77) Perithecia elevated, often craterous. juga forming a reticulate pattern (Fig. 34); thallus opaque dp,', translucent wet

Verrucaria amphibia

Figure 34.— a. Vertical section of perithecium. Scale is 1 mm. b. Thallus of Ver-
rucaria amphibia. (Specimen illustrated is thicker and stored longer than that in
Fig. 39b.) Scale is 1 mm.

19 (17) Juga appearing as pegs prottuding from thallus and also to some extent on penthecia; thallus nmose-areolate.
usually blackish-brown and opaque (wet or dry); perithecia often large, 0.3-0.7 mm diameter, immersed or
elevated, gently sloped to nsing abruptly, sometimes depressed on top (Fig. 35); spores 10-20 x 7-lOfim Verrucaria maura

Figure 35.— a. Vertical sections of perithecium. Scale is 0.1
mm. b. Thallus i>f Verrucaria maura. Scale is 1 mm.

12

19 (/7) Juga raised or immersed, appearing as usually irregular and often branched ridges often merging with or continuing
over penthecia, if immersed appearing as black spots when wet; thallus blackish-brown, becommg brown to
amber and translucent when wet; perithecia spreading and irregular at base, up to 0.3 min diameter (Fig. 36);
spores 8-9 x 4.5-7/im Verrucaria erichsenii

Figure 36.— a. Vertical section of perithe-
cium. Scale is 0.1 mm. b. Thallus of Verruca-
ria erichsenii. Scale is 1 mm.

20 (/6) Perithecia immersed, spores 9-12 a 5-6;(m; juga appearing as long sharp black ridges on borders of thallus, some-
times extending across thallus, fissures occasionally develop along juga and simulate areolation; thallus usually
green to cream colored (Fig. 37) Verrucaria degelii

Figure 37.— a. Vertical section nf perithecium. Scale is 0.1 mm. b. Thallus of Verrucaria degelii

Scale is 1 mm.

20 {16) Perithecia raised and prominent (Fig. 38) 21

Figure 38.— Thallus with prominent raised
perithecia.

21 {20) Juga inclosing lenticular areas giving appearance of npples on water, continuing on pentheciutii to form reticular

pattern. Perithecium elevated, often craterous (Fig. 39); spores 9-20 x 4-7.5;tni Verrucaria amphibia

Figure 39. — a. Vertical sectit)n of perithecium. Scale is 1 mm. b. Thallus of Verrucaria amphibia. (Thai
us thinner than Fig. 34b. Cracks developed in storage.) Scale is 1 mm.

21 [20) Juga scattered, not inclosing lenticular areas 22

13

22 (2/) Juga appeanng as pegs to usually short ndges, usually straight but sometimes crescent shaped, sometimes ccia-
iescing with perithecium at base; thallus usually dark olive green but lighter and with fewer juga in shade; pen-

thecia hemispherical, often shiny (Fig. 40); spores 6-11 x 3-5^m

. Vernwana d it mar sic a

Figure 40, —a. Vertical section or perithecium. Scale is 0.1 mm. b. Thallus of Verrucaria ditmarsica

Scale is I mm.

22 {21) Juga broader, irregular, and sometimes branched (Fig. 41)

.23

Figure 41.— Thallus with broad, sometimes branched
juga. Scale is 1 mm.

23 {22) Juga conspicuous, highly irregular, frequently branched, raised or immersed and appearing as conspicuous black
spots in wet thallus. often merged with penthecia; wet thallus usuall) brown to amber and translucent (Fig. 42):
penthecia up to 0.3 mm diameter, spores 8-9 x 4.5-7^m Verrucaha crichscnii

Figure 42. — a. Vertical section of
perithecium. Scale is II. 1 mm. h.
Thallus of Verrucaria erichsenii.
Scale is I mm.

14

23 (22) Juga prominent, black, and shiny, much thicker than in V erichsenii. frequently broadened into thick irregular
plates, especially at thallus margin; thallus usually grass green but darker in the sun and depigments rapidly in
shade or in storage; perithecia typically heniisphencal to globular with flattened tops but may become quite
irregular, angular, or dissected (Fig. 4.1); spores 8-10 x 4-6fim Vernicaihi siriciluhi

Figure 43.— a. Vertical section of
perithecium. Scale is 0.1 mm. b.
Thallus of Verrucaria strialula.
Scale is 1 mm.

ANNOTATED LIST OF SPECIES

Where distributions were derived from the literature only,
author and date are given. All other specimens were examined by
the author Where such specimens were collected by others, the col-
lectors name is given followed by the identification of the herbar-
ium retaining the specimen. The remaining distnbutions are from
the authors collections.

Genus Arthopyrenia

Anhopyrenia halodytes (Nyl.) Am. Ber Bayer Bot. Ges. 1:122.
1891. Verrucaria luiloihles >iy\. Mem. Soc. Sci. Nat. Cherbourg
5:212. 1857.

DESCRIPTION: Thallus epilithic (siliceous rock), endolithic
(calcareous rock and shells), yellowish if epilithic, greyish or
blackish-brown if endolithic. Perithecium almost wholly immersed
in substrate or sessile with black hemispherical to broadly conical
involucrellum, 0.15 to 0.5 mm diameter, excipulum colorless to
pale brown. Spores eight perascus. ovoid, two-celled, one cell usu-
ally broader than the other. 9-20 x 4-7.5;(m.

Sw inscow (1965) considered this to be the only littoral species
of the genus and treated A. sublitoralis (Leight.) Am.. A. foveolaia
A. L. Sm. and A. i^yaleaoidea Knowles as synonyms and reported
a range of 10-20 ■; 5-10/im in spore size. Richard C. Harris ( 1975)
treats this species as co. .a PyrenocoUemu.

Anhopyrenia halodyles is often associated with shells or
calcareous rocks but may also be found on siliceous rock.

GENERAL DISTRIBUTION: Worldwide on marine shores
(Santesson 1939:52-63).

NORTHEASTERN AMERICAN DISTRIBUTION: NEW
JERSEY: Cape May Co. CONNECTICUT: New London Co.
RHODE ISLAND: Newport Co. MASSACHUSETTS: Barnstable

Co.. Plymouth Co., Essex Co. MAINE: Cumberiand Co.. Sagada-
hoc Co., Hancock Co.. Washington Co. NOVA SCOTIA: Yar-
mouth Co., Digby Co., Halifax Co., Victoria Co., Cape Breton
Co. NEWFOUNDLAND: West Coast Section, Northern Penin-
sula Section. East Coast Section. Avalon Section.

Genus Caloplaca

All of the littoral species of Caloplaca collected belong to the
section Gasparrinia.

Caloplaca marina (Wedd.) DuRietz, Method. Grund. Modern.
Pflanzensoziol. 170. 1921. Lecanora marina Wedd. Mem. Soc.
Sci. Nat. Sherbourg 19:275. 1875.

DESCRIPTION: Thallus orange yellow to red orange, citnne in
shade, orbicular, irregular, or subeffuse, small convex lobes contig-
uous or. in center of thallus, minutely granular or tuberculate, whit-
ish prothallus sometimes visible. Apothecia reddish-orange,
0.5-1.0 mm diameter, plane to convex, margins entire or crenulate.
Spores ellipsoid, polarilocular, isthmus about 0.3 length of spore,
10-14 X 5-7,im.

Wade (1965) found this species often associated with Caloplaca
tballincola and Verrucaria maura in the British Isles. Caloplaca
ihaliincola has not been collected, however, in the littoral zone of
North Amenca. The white hypothallus is seen in young growth
between the small subconvex lobules.

GENERAL DISTRIBUTION: Norway, Sweden. Poland (Nor-
din 1972). Finland (Rasanen 1927). Germany (Enchsen 1957),
Novaya Zemyla (Lynge 1928), England (Ferry and Sheard 1969),
Wales (Fletcher 1973b), France (Werner MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberland Co., Hancock Co. NOVA SCOTIA: Yarmouth Co.,
Digby Co., Shelburne Co., Halifax Co. NEWFOUNDLAND:
Northern Peninsula Section.

15

Cidoplaca micmlhalliiw (Wedd.) Zahlbr. Cat. Lich. Univ. 7:247.
1931. Lecanora microlhallind Wedd. Mem. Soc. Sci. Nat. Cher-
bourg 19:276. 1875.

DESCRIPTION: Thallus citrine. 1-3 mm diameter, compnsedof
scattered lobes on the peripher>. scattered granules toward center,
forming patches on rocks, lobes convex. 1-2 times as long as
broad, tending toward granules near center of patches, Apothecia
yellow to pale orange. 0. 15-0.8 mm diameter, margins crenulate or
entire. Spores oblong-ellipsoid, polarilocular. isthmus ca. 0.3 the
length of the spores, 10-18 x 5-8/4m.

Caloplaca microthallimi is distinguished by its minute granular
thallus forming patches up to 3 mm diameter and by its citrine
color Wade ( 1965) found it often associated with Verruairia muuni
in the British Isles. This also holds true for North America. This
species is not previously reported from North America. In Europe
this species is not commonly reported from the littoral zone.

GENERAL DISTRIBUTION: Norway, Sweden, Finland. Den-
mark (Nordin 1972).

NORTHEASTERN AMERICAN DISTRIBUTION: NOVA
SCOTIA: Digby Co.. Halifax Co., Cape Breton Co. NEW-
FOUNDLAND: West Coast Section. Northern Peninsula Section.

Culopliua scopiilaris (Nyl.) Lett. Hedwegia 52:242. 1912.
Lecanom scopulahs Nyl. Flora 66: 105. 1883.

DESCRIPTION: Thallus yellow orange to deep orange, radiate.
up to 1 .5 cm diameter, lobes narrow, 0.25-0.3 x 0.5-2.0 mm, con-
vex, apices crenulate or branched, center of thallus usually thickly
covered with apothecia. Apothecia orange, 0.5-0.75 mm diameter,
margins entire. Spores ellipsoid, polarilocular, 10-15 x 5-7/im.
isthmus ca. 0.3 the length of the spore.

Caloplaca scopiilaris is unique among the littoral Caloplacae due
to its radiating thallus. Wade (1965) indicated that this species
resembles a small form of C. heppiana or C. thallincola but is dis-
tinquished by ellipsoid spores and the submoniliform nature of the
upper parts of the paraphyses.

GENERAL DISTRIBUTION: Novaya Zemlya (Lynge 1928),
Norway, Sweden, Denmark (Nordin 1972), Finland (Rasanen
1927), Germany (Enchsen 1957), lapan (Ny lander 1890).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberland Co., Hancock Co. NOVA SCOTIA: Halifax Co., Vic-
toria Co. NEWFOUNDLAND: West Coast Section, Avalon Sec-
tion.

Caloplaca vcrniciilifera (Vain.) Zahlbr Cat. Lich. Univ.
7:272.1931. Placodium verniculifera Vaimo in Med. om Gron.
30:313. 1905.

DESCRIPTION: Thallus orange or yellow, up to 3 cm diameter,
ordispersed, lobes 3-5 mm long, convex, centerof thallus areolate.
granular, isidia globose, spores oblong-ellipsoid, polarilocular,
isthmus ca. 0.3 length of spore, 9-14 x 4-6/im.

Specimens of this species, previously unreported from North
America, were identified for the author by J. Poelt. It is easily con-
fused with Caloplaca granulosa which also has globose isidia but

has smaller rosettes of 1-1.5 cm and shorter lobes of about 2 mm
(Poelt 1969).

GENERAL DISTRIBUTION: Norway, Sweden. Finland, and
Siberia (Nordin 1972).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Sagadahoc Co., Hancock Co. NEWFOUNDLAND: West Coast
Section.

Genus Lecanora

Lecanora graiuii Magn. Ann. Cryptog. Exot. 5(1):21. 1932.

DESCRIPTION: Thallus of irregular whitish-grey large gran-
ules. KOHh- yellow, medulla with small crystals. Apothecia con-
cave to plane, thalline margin regular to crenulate; hymenium dark
reddish-brown, I + dark blue; epithecium with small crystals,
insoluble in KOH. Spores eight per ascus, oblong-ovoid, 11-18 x
6-8.5/(111.

Littleof this species was collected since it extends down only into
the upper limits of the littoral zone. The scant material collected
conforms to the original niatenal described from a log on a sea
beach by Magnusson (1932).

GENERAL DISTRIBUTION: Washington (Magnuson 1932),
British Columbia (Brodo MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Sagadahoc Co. NEW YORK: Suffolk Co. (Latham MSC). NOVA
SCOTIA: Shelburne Co. NEWFOUNDLAND: West Coast Sec-
tion.

Genus Lichina

Lichina confinis (O. Mull.) Ag. Sp. Algar. 1:105. 1821. Lichen
ConfinisO. Mull Icon. PI. Daniaw 5:5. 1872.

DESCRIPTION: Thallus fmticose. branches rounded in cross
section, blackish-brown, phycobiont Calothrix. Ascocarps termi-
nal, globose, perithecoid, small ostioles. Spores colorless, simple
ovoid. 15-24 x 12-15/im.

Lichina confinis. appearing as black tufts in the upper limits of
the littoral zone, is the only fmticose lichen found in the area
included in this treatment.

GENERAL DISTRIBUTION: England (Ferry and Sheard
1969). Wales (Fletcher 1973b). Finland (Rasanen 1927), Germany
(Degelius 1939). Italy (Jatta 1909-191 1). Norway (Havaas MSC),
Sweden (Degalius MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MASSA-
CHUSETTS: Essex Co. (Tuckerman 1882). NOVA SCOTIA:
Cape Breton Co. (Lamb 1954). NEWFOUNDLAND: East Coast
Section. Northern Peninsula Section. West Coast Section.

Genus Stigmidium

Stigmidiiim marinum (Deak.) Swins. Lichenologist 3:55. 1965.
Sagedoa marina Dcak. Ann. Mag. Nat. Hist. I. 12:40, pi. 4, f
13. 1954.

16

DESCRIPTION: Parasymbiont on littoral Verrucariae, espe-
cially V mucosa and V microspora. lacking a visible thallus of its
own. Perithecia usually resembling that of host lichen, totally
immersed on V. mucosa and hemispherical on V microsponi. Invo-
lucrellum black with pigment extending into hyaline excipulum.
Spores eight per ascus. two-celled with each cell often divided by
pseudosepta. uppercell usually slightly widerthan the lower, 10-15
X 3.5-5/im.

The perithecia of Stigmidium are easily confused with that of the
host. Sligmidium. however, has two-celled spores of 10-15 x 3.5-
5/im. Swinscow (1965) reported spores 10-15 x 4-6/im. The com-
bination Stigmidium marinum was made by Swinscow when he
united Arlhopyrenia marina (Deak ) A. L. Sm and A. leptotera
(Nyl. ) Am. Neither of these species appear in the North American
checklist (Hale and Culberson 1970) nor did Swinscow (1965)
report a specimen from North America. This, therefore, represents
a new record for North America.

GENERAL DISTRIBUTION: Germany, Finland, Ireland, Jer-
sey. England (Swinscow 1965).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberland Co. , Hancock Co. NEW JERSEY: Ocean Co. NEW-
FOUNDLAND: Avalon Section.

Genus Verntcaria

The species of this genus form crusts which may be contmuous or
form scattered patches, rimose areolate or divided into discrete are-
oles or lobes. The thalli vary in thickness from 20 to 500/jm, and
may be almost completely transparent to entirely opaque. In some
species, the transparency of the thallus may be enhanced by wet-
ting, while others remain opaque when wet. The thalli may be
smooth or roughened by dark points or ridges. The ridges, called
juga (Santesson 1939), vary in length, width, and thickness. The
exact ongin and ontogeny of the juga is not known but it appears
that they originate at or just below the surface of the thallus. In the
author's experience, they do not extend to the substrate except at the
edges of the areoles. Often a distinction is made in keys between
point and ridge forms of the juga, but it is common that the points
are merely peaks on inconspicuous or hidden ridges.

The perithecia of all of the littoral Verrucariae, with which the
author has worked, have a dark involucrellum which may spread
widely or be closely appressed to the excipulum. This darkening
may extend into the lower excipulum, terminating a short distance
below the involucrellum proper or it may not extend into the lower
excipulum. The darkening may occur intermittently or continu-
ously throughout the lower excipulum. This darkening is given
considerable taxonomic importance by Servit ( 1954) and by Ench-
sen (1957) but Swin.scow ( 1965) discounted this importance on the
grounds that it is highly variable within a species. The author con-
curs with Swinscow in this view. Perithecial size is not, in itself,
very useful in distinguishing species. Perithecium diameters vary
generally between 0.05 and 0.7 mm and these dimensions overlap
considerably between Verrucaria species. Although vertical sec-
tions are illustrated in the key. none is claimed to be typical for then;
is much variation in many, if not all species.

Santesson (1939) has noted the high variability of spore size in
littoral pyrcnolichens and suggested that they should not be given
great taxonomic importance. All spores in yernicariti are simple
and hyaline. For the most part, the spores are thin walled and of
similar size, occurring eight per ascus. Verrucaria inlernigrescens

and V silicicola are the notable exceptions to the above in north-
eastern North America. The spores of V. inlernigrescens are thick
walled and those of V. silicicola are unusually large. The remainder
are ovoid to reniform and exhibit overiapping size ranges between
species,

Verrucaria amphibia R. Clem. Ess. Var Veg. Andalucia 299.
1807.

DESCRIPTION: Thallus continuous to rimose areolate. black
when dry. translucent green to amber when wet, with irregular
black ndges often enclosing lenticular areas 50-l(X)/im thick.
Perithecia large (13-16 mm), prominently elevated, often flattened
or concave on top. vertical and horizontal ridges forming reticular
pattern, lenticular areas often enclosed by ridges at base; excipu-
lum hyaline to partly darkened below. Spores hyaline, ovoid. 7-19
X 4-7. 5^m.

This species might readily be confused with V. maura. Both spe-
cies have dark thalli with juga forming pegs or ridges and with
robust penthecia. However, they are easily distinguished on closer
examination. Whereas V. maura tends to have points raised above
the thallus, V. amphibia tends toward ridges. The ridges tend to
enclose lenticular areas on a dry thallus giving the impression of
ripples on water When wet. V. amphibia becomes more translucent
and light colored against which the black ridges become contrasted.
The thallus of V. maura remains dark when wet except on very thin
juvenile specimens. The thallus of V. amphibia tends to be continu-
ous whereas that of V. maura tends to be rimose areolate but in nei-
ther case are the characters absolute. Areolation appears to develop
and increase with age in V. amphibia. Verrucaria amphibia pro-
duces large, steep sided perithecia often strongly depressed on top.
a form also produced in some instances by V maura. Such perithe-
cia of V. amphibia have two characteristics not demonstrated by V.
maura: whereas V. maura may have many fine points or pegs cov-
ering the perithecium. the design on V amphibia is a reticulum of
vertical and horizontal ridges. Also, where the V amphibia perithe-
cium meets the thallus, the ridges tend to enclose areas of ovoid
shape and appear light and translucent when wet (see the Key).

Verrucaria amphibia is little discussed in literature. Santesson-
says that it is not known from Scandinavia, although it occurs in
England. Ferry and Sheard (1969) included V amphibia in their key
but did not deal with it in detail . The author first became acquainted
with It by examining herbanum specimens and his measurements
should not be regarded as limits due to the small amount of material
examined. There is no prior report of this species from North
Amenca.

GENERAL DISTRIBUTION: England (Ferty and Sheard
1969). Wales (Fletcher 197.3a). Germany (Ullnch MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: NEW-
FOUNDLAND: East Coast Section. Avalon Section.

Verrucaria ceuthocarpaWMenb. in Ach. Suppl. Meth. Lich. 22.
1803.

DESCRIPTION: Thallus brown, rimose areolate, glabrous,
forming discrete areoles or lobes, ridges absent, perithecia sub-

-R. SanlcsMin. Dirctlur. S\sienuilic Insiiiute of Oslo. Oslo. Nor^^.;l\. pcrs
commun November 146(1.

17

merged. 0. 1-0. 15 mm wide, involucrellum usually brown, excipu-
lum hyaline to dark below, spores hyaline, ovoid 8-10 x 5-8fim.

Verrucaha ceuihocarpa is easily confused with Verrucariu tlege-
lii when either is modified by environment or when its structure is
obscured by epiphytic algae. The clefts of V. cethocarpa may
become darkened and. on cursory examination, may resemble the
dark ridged border of areoles of V. degetii. Careful examination is
needed to observe the absence of ridges. Conversely sometimes the
ridges are scant on a thallus of V. degelii and it may by mistaken for
that of V. ceulhncarpu .

Perhaps the greatest obstacle to correct identification is obscur-
ing of surface detail by epiphytic algae. Portions of questionable
thalli should be flooded with water to soften and expand the algae.
If details are still obscured, the flooded portion can be scrubbed
with a small water color brush. True ndges are fairly resistant to
such gentle abrasion and will persist until the thallus becomes too
.softened by the water to maintain its integrity.

According to Santessons key (1939), the pnme diagnostic feature of
k' degelii vs. V ceuthocarpa is the presence of ridges on V. degelii.

The author finds that two other features are quite useful. The best
of the two is tendency for tlatness to concavity between ndges of an
areole of V. degelii compared with the convexness of areoles and
branches thereof in V ceiilhocarpa. Verntcaria ceulhoearpii is truly
glabrous. Further, from the limited material in the authors three
North American collections it also appears that the involucrellum of
V ceuthocarjxi tends to be brown whereas that of V. degelii is dis-
tinctly black.

GENERAL DISTRIBUTION: Northern Europe. Spitsbergen,
Bear Island, Novaya Zemlya, Sibena. Bering Straits. Greenland,
Kerguelen, Antarctica (Lamb 1953); Washington U.S.A. (collected
by Fink and determined as Verruearici nuiiira by Zahlbruckner
MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: NEW-
FOUNDLAND: East Coast Section.

underlying thallus and reduce the growth of ndges. On the other
hand, a growth of dark algae on V ceuthoearpa, especially when dry,
may obscure the color and glabrous nature of the thallus and give the
illusion of ndges on the edge of areoles. One cannot (wer-emphasize
the importance of detecting and removing such epiphytic algae!

The thallus of V. degelii varies greatly in thickness and transpar-
ency depending on age or extent of development. It is almost always
areolate. with only one known exception. In that case, a rather large
area of smooth, continuous thallus was surrounded by a raised black
ndge and a crack, also bordered by a black raised ridge, extended
inward t(iward the center At the edges of this patch of thallus was
more thallus divided into typical discrete areoles with similar black
borders. More commonly a young thallus is thin, nearly transparent,
smooth and brown to tan. It is areolate and is often divided into lobes
by a dendntic pattern of grooves. In such a thallus. the grooves are
usually hyaline.

The ontogeny of these lichens has not been studied and must be
inferred from observations of what could as easily be the results of
conditionsof growth as of aging. It appears, however, that older thalli
thicken and develop discrete areoles, usually with black borders and
frequently with black lines connecting them with the pentheciuni.
These areas of blackness then seem to proliferate under some condi-
tions to produce unelevated patterns or raised juga. Sometimes the
black area extends beneath the perithecium and sometimes it docs
not.

The penthecium is commonly immersed, with or without raised
ndges aniund the ostiole. At times there is a convex area raised around
the o.stiole. The author has seen prominently raised convex perithecia
on the same thallus as sunken perithecia. The degree of penthecial
elevation appears to be a vanable character of the species.

Spores are ovoid but sometimes pointed when young. A clear spot
commonly, although not always, appears in the center of the younger
spores. Sanlesson (1939) reported the range of spore sizes for Verru-
cariu degelii as 10-13 '■ 5-6/im. which is consistent with that
observed by the author

GENERAL DISTRIBUTION: NORWAY (Santesson 1939).

Vemicaria degelii R. Saiit. Ark. Bot. 29A (10)31. 19.^9.

DESCRIPTION: Thallus rimose, forming lobes or discrete are-
oles, amber, brown or black when dry, amber to brown when wet,
prominent black ridges bordering areoles and often perithecia. thin to
thick (l(K)-320 mm). Perithecia sunken to prominently raised, 0.1-
0.25 mm diameter, excipukim hyaline. Spores hyaline, ovoid 9-13 x
5-6/(m.

Though published in 1*^39. this species was known only from Nor-
way until the author's collections, thus there are few specimens and
little study of ecology upon which to base conclusions about environ-
mental intlucnces on variations. It is reasonable to assume, however,
that the nature of variations is consistent with that seen in other Vemi-
canae. If so. color wduUl he lighter in shade forms than sun lorms
and ndges would be fewer and thinner in shade forms than on sun
forms. This kind of variation is probably the cause of confusion o\
some specimens of V degelii with V ceiilhocarpa . The identity of a
robust .specimen of V degelii is beautifully clear. However, the ridges
may become scant and the specimen strongly resembles the morpho-
lype of V ceutluKurjui with darkened clefts between areoles. A sec-
ond source of conftision is the cpiphvtic growth of algae on thalli of
cither \' degelii or 1.' ceiithtHiirpa- A growth of algae on I,' degelii
can obscure ndges and produce the glabrous appearance of I.' cemlu>-
carpu. Such a growth can also produce a shade condition for the

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Hancock Co.. Washington Co. MASSACHUSETTS: Essex Co.
NOVA SCOTIA: Yarmouth Co., Digby Co.. Halifax Co. NEW-
FOUNDLAND: West Coast Section, Northern Peninsula Section,
East Coast Section, Avalon Section,

I'erntcana diliiiarsua Erichs. Schriften Naturwiss. Vereins
Schleswig-Holstein 22:90. 1937.

DESCRIPTION: Thallus entire, olivaceous, with pegs to short
ridges, thin (20-50^m), opaque to translucent when dry, more
translucent when wet. Pcnthecia slightly dome-shaped to globose
(usually henusphencal). 0.1-0,25 nnii diameter. usualK shiny;
excipulum hyaline to dark below. Spores o\i>id to reniform. color-
less, 6-1 1 X 3.5-6^(111.

The thallus of Verntcaria ditinarsiva is usually olivaceous, but
the darkness varies with light exposure. In bnght light the thallus is
dark olive and if grown in the shade it is a very light olive green. It
usually is rather transparent to translucent. The transparency is
increased by wetting upon examination. Short, mostly straight
ndges are charactenstic on the thallus. These become reduced in
size and frequency in the shade modification. In this reduced condi-
tion it is easily confu.sed with the shade modification of I,' erich-

18

senii and especially with V siriatula whose color is also similar
Ridges often merge with the peritheeia at the base.

The penthecia are convex to hemispherical and usually shiny.
Darkness of the lower excipulum is a highly variable factor Gener-
ally some carbonaceous granules lend to extend below the mvolu-
crcllum and may continue somewhat throughout the lower
involuerellum (sec the key). However, the species should not be
charactenzed as having an involuerellum that is dark below, a view
adopted by Erichsen ( 1957).

GENERAL DISTRIBUTION: Germany (Erichsen 1957), Nor-
way (Santesson MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberiand Co., Sagadahoc Co., Hancock Co. RHODE
ISLAND: Newport Co. MASSACHUSETTS: Plymouth Co.,
Essex Co. NEW HAMPSHIRE: Rockmgham Co. NOVA SCO-
TIA: Yarmouth Co., Digby Co.. Shelburne Co., Halifax Co., Vic-
tona Co. NEWFOUNDLAND: West Coast Section. Northern
Peninsula Section, East Coast Section. Not previously reported
from North Amenca.

Vcrnicaria crichsenii Zsch. in Erichs. Verb. Bot, Vereins Prov.
Brandenburg 70: 192. 1928.

DESCRIPTION: Thallus entire or areolate, black to blackish-
brown dry, amber and more translucent when wet, roughened by
rows of pegs or ndges; ridges often branched, sometimes sub-
merged in thallus and visible only when wet; thallus 30-70/jni
thick. Penthecia elevated, conical to hemispherical, spreading in
irregular pattern at base, often with pegs or ridges as on thallus.
0. 1-0.3 mm diameter, excipulum hyalme below. Spores ovoid, col-
orless. 8-9 X 4.5-7fim.

Vernicaria erichsenii demonstrates considerable thallus varia-
tion. It is usually considered to be cleariy rimose or areolate. Typi-
cally this is so but it is not uncommon to find a thallus of V.
erichsenii continuous in an especially wet environment, where the
thallus becomes thick and gelatinous. When dry the thallus is usu-
ally blackish-brown to black, A grey morphotype with a texture
like that of graphite was collected in the Narragan.sett Bay area.
When moistened during examination the thallus increased in trans-
parency, one of the best tests of questionable thalli. When wetted
they reveal a pattern of black markings typical of the pattern of
ridges usually seen above the surface. The thallus of V. erichsenii
typically has short, often branched irregular ridges or rows of
points, frequently confluent with or continuing up over the penthe-
cia. Ridges are longer than wide and rarely very high. In extreme
circumstances ridges become higher and sharper forming cusps and
aretes. In other extremes they may be immersed within the thallus
to be revealed only by wetting.

The most consistent character of the penthecium appears to be
the irregular spreading base which seems most pronounced when
viewed from above a wet thallus or when the thallus has been
peeled from the rock and placed on a slide and illuminated from
below. The diameters of penthecia tend to fall between 0. 1 and 0.3
mm.

Zschacke (1934) and Enchsen ( 1957) both list a spore range of 8-
12 X 5-7;(m which agrees with the author's observations.

GENERAL DISTRIBUTION: Germany (Enchsen 1957), Bnt-
ish Columbia (Ohls.son MSC), Sweden (Santesson MSC). Nonvay

(Santesson MSC). Wales (Brodo MSC). Scotland (Brodo MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberiand Co.. Sagadahoc Co.. Hancock Co.. Lincoln Co.
(Memll MSC). Washington Co. MASSACHUSETTS: Essex Co..
Bristol Co. (Willey MSC). Plymouth Co., Barnstable Co. RHODE
ISLAND: Newport Co., Washington Co. CONNECTICUT; New
London Co. NEW HAMPSHIRE: Rockingham Co. NOVA SCO-
TIA: Yarmouth Co., Digby Co., Shelburne Co. . Halifax Co., Vic-
toria Co.. Cape Breton Co. NEWFOUNDLAND: West Coast
Section, Northern Peninsula Section, Avalon Section,

Verrucarui infcrnif^rescens (Nyl.) Enchs. Vern. Bot. Vereins Prov.
Brandenburg 70:193. 1929. Vernicaria aethiobola van Inlerni-
f^rescens Nyl. in Brenner. Meddeland. Soc. Fauna Fl. Fenn
13:125. 1886.

DESCRIPTION: Thallus of brown areoles connected by darker
brown prothallus. Peritheeia brown, convex, 0.2-0.34 mm diame-
ter, excipulum hyaline or intermittently darkened below. Spores
hyaline, thick walled, fusiform. (11.5) 15-23 (27) x (4) 6-8
(I4)fim.

The brown prothallus. thallus. and penthecia are distinctive. The
thallus is thin by comparison with other areolate forms such as V
maiira and V ceiilhocarpa. The spores are distinctive, being fusi-
form and thick walled. Erichsen ( 1957) reported a spore range of
15-27 X l-\2nm. As indicated above, spores were found in the
North American collections as large as reported by Enchsen but
they were bloated and of atypical shape. Most spores seen were in
the range of 15-23 ^ 6-8;(m. Erichsen (1957) listed this species as
both littoral and nonlittoral. therefore, it may be assumed to be
facultative in its habits. This species was collected at only one
locality in North America and this is the first record from North
America.

GENERAL DISTRIBUTION: Germany (Enchsen 1957).

NORTHEASTERN AMERICAN DISTRIBUTION: NEW-
FOUNDLAND: Avalon Section.

Vernicaria maiira Wahlenb. in Ach. Suppl. Meth. Lich. 19. 1803.

DESCRIPTION: Thallus dark, usually black to brownish-black,
sometimes green, opaque (wet or dry), rimose areolate. usually
with small black pegs or points (sometimes obscured by epithallic
algae), thin to thick (75-300 ^mi). Peritheeia small to large (usually
large) 0.3-0.7 mm diameter, sunken to prominently raised tops
rounded to concave, may be covered visibly by pegs. e,xcipulum
entirely black below. Spores ovoid, hyaline 10-20 x 7-10;im.

Vernicaria nuiitra is probably one of the most widely used names
among the littoral lichens. This often leads to the report of the spe-
cies based on the most cursory examination. The species is highly
variable and its variation has probably been best understood and
described by Santesson ( 1939. footnote 2). The author can add little
to the descnption of variation provided by Santesson (footnote 2)
and would like here to merely record his concurrence and quote
Santesson's description (the material quoted was provided for the
author in Swedish by Santesson) of typical V nuiiira as ; " 1 . Indis-
tinct or dark prothallus. 2. Definite black thallus. 3. areolate thal-
lus. 4. thallus surface with distinct but little conspicuous points, 5.

19

rounded conical conspicuous perithecia. 6. rounded, not impressed
perithecia tops, and 7. spores with a width of 7-10 mic. (length 10-
19 or at most 20 mic.)"

Santesson continues by descnbing seven deviations with which
he associates names of varieties or what he considers to be syno-
nyms:

■"1. With while or light broun clear prothallus. 'V' -scluickccinii
Erichs."

2. With green-gray or greenish black thallus. 'van fiimosocine-
rea Vain.'

3. With connected or tnflingly fissured thallus. 'V. scotinu
Wedd.'

4. Thallus surface with many strongly conspicuous pegs. 'V.
iinutiiui Wg."

5. a. With perithecia almost entirely sunken into the thallus. 'V

malmei Serv.'
b. With hemispherical to nearly sphencal penthecia. "var
pniminuUi Vain."

6. With clearly impressed, frequently somewhat irregular
perithecium tops. 'V. trachinodes Norm.' 'V. hayrcnti
Erichs.". 'var araclinoides Vain", "f. evoluta Th. Fr"

(?7. Spores with a width of 1 1-15 mic. ' V. finnnuirkicti Zsch.' )."

The author has seen all of these forms in North Amenca except '" V.
finnmarkica." A common variation results from frequent washing
with water from rain or waves which causes the thallus color and
texture to become obscured by epithallic algae which produce a
smooth greenish surface.

GENERAL DISTRIBUTION; Novaya Zemyla (Lynge 1928).
Finland (Rasanen 1927). Germany (Erichsen 1957). Spain to Fin-
land (Degelius 1935). Italy (Jatta 1909-1911), India (Awasthi
1965), Japan (Nylander 1890). Greenland. Iceland, Bear Island.
Spitsbergen. Siberia. Bering Strait. Fuegia. Patagonia. Chile.
Falkland Islands, and New Zealand (Lamb 1953). Norway (Santes-
son MSC). Sweden (Santesson MSC), Wales (Fletcher 1973a).
England (Ferry and Sheard 1969). British Columbia (Brodo
MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
(Plitt USA). NOVA SCOTIA: Yarmouth Co.. Digby Co..
Shelburne Co.. Halifax Co.. Victona Co.. Cape Breton Co. NEW-
FOUNDLAND: West Coast Section. Northern Peninsula Section.
East Coast Section, Avalon Section. NEW BRUNSWICK:
Campobello

wet. Erichsen (1957) recognized a var laetevirens as characterized
by a leek green thallus which is less transparent. The author has
also observed this in material from the southern part of its range in
North America as well as a dark grey thallus in the same area. The
dark grey thallus has a color and texture resembling graphite used in
pencils. Upon wetting, the dark color disappears.

The penthecia are brown to black and may be shiny or dull. Their
shape may range from that of a slightly arched dome to more nearly
hemispherical and pointed. The diameter of the perithecia is
reported as 0.15-0.25 mm (Lamb 1953). 0.1-0.3 mm (Santesson
footnote 2), 0.2-0.3 mm (Zschacke 1925). and 0.2-0.3 mm ( Erich-
sen 1957). Erich.sen ( 1957) also recognized xar friesiaca as having
a smaller perithecia of only 0.1-1.15 mm diameter The author
measured 230 penthecia (10 each from 23 specimens) and found a
range of 0.08-0.28 mm except for a single collection in which the
range was 0. 16-0.48 mm with 4 out of 10 exceeding 0.3 mm. In no
other collection did the diameter exceed 0.3 mm. Clusters of small
or larger perithecia may be found and due to the normal patchy
appearance of the thalli. one may be tempted to consider such
patches as distinct species. Penthecium size should not be consid-
ered of taxonomic significance in this species.

Spores of V microspora are thin walled and will change shape
somewhat with osmotic changes. Generally they are ovoid to slightly
reniform. The species name infers smallness of spores, but this is not
noticeably different from I' ditnuirsua and V sirUituUi. Spore sizes
have been reported as 7-12 x 4-5//m (Lamb 1953), 7-11 x 5-7;(ni
(Zschacke 1925), and 7-11 x 4-5fim (Erichsen 1957). Enchsen
(1957) also recognized var mucosula Sands, with spores 5-9 x 4-
9;im, The author's spore measurements give a range of 6-11 x 3-
5;(m.

GENERAL DISTRIBUTION: Germany (Erichsen 1957),
England (Ferry and Sheard 1969), Greenland, Japan, Chile, Ant-
arctica (Lamb 1953), Wales (Fletcher 1973a), Norway (Santesson
MSC). Scotland (Brodo MSC). Australia (Willis MICH).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Cumberland Co., Sagadahoc Co.. Hancock Co. MASSACHU-
SETTS: Essex Co.. Plymouth Co.. Barnstable Co. RHODE
ISLAND: Newport Co. CONNECTICUT: New London Co. NEW
JERSEY: Ocean Co.. Cape May Co. NOVA SCOTIA: Colchester
Co., Cumberland Co., Yarmouth Co.. Digby Co.. Shelburne Co.,
Victona Co. NEWFOUNDLAND: West Coast Section. Northern
Peninsula Section. East Coast Section.

Verrucanannuosa ^'Mcnh. in.Ach. Suppl. Mcth. Lich. 23. 1803.

Vernicdiut inurDsponi Nyl. Ann. Sci. Nat. Bot. IV. 3: 175. 1855.

DESCRIPTION: Thallus usually brown to amber, sometimes
green or grey, smooth thin. 20-30 iim continuous or in patches but
never areolate. translucent when dry, more transparent when wet.
Penthecia dome-shaped (slightly arched) to hemisphencal, some-
times pointed. 0.05-0,3 mm diameter, excipulum entirely hyaline
below. Spores hyaline, reniform to ovoid. 6-1 1 x 3-5/im.

Vernuciria microspora is characterized by a thin. normalK
brown to amber thallus which is cither continuous or in scattered
patches but never areolate. The thallus is devoid of pegs or ndges
but may contain artifacts that might superticially resemble pegs or
ridges. In some instances tests ha\c proven these to be perilhecial
remains. The thallus becomes translucent to quite transparent when

DESCRIPTION: Thallus grass green to blackish-green, smooth
and tough, continuous, often with necral cracks in herbanum speci-
mens, 130-150 ;im thick, usually opaque (wet ordry); juga absent;
prothallus. is present, whitish. Penthecia submerged to slightly
raised, sometimes with prominent ostioles surrounded by chimney,
excipulum clear below, involucrellum 0.05-0.2 mm diameter
Spores simple, coloriess. usually ovoid. 8-11 ^ 4-5 /im.

Thallus of this species is easily recognized in the field or in the
herbarium. In old herbanum specimens it is brown. Upon drying it
develops cracks that arc stnctly necral artifacts not to be confused
w ith the nmose areolation typical of V. maiira. A whitish prothallus
is often seen around a thallus on smooth rock.

Perithecia arc normall) flush to slightly raised or sunken. When
ripe, the nnolucrellum tends to cvaginate somewhat presenting a

20

conspicuous ostiole with a short chimney around it. One often sees
whitish pits in the thallus which are the remains of old penthecia
divested of their involucrella.

The thallus of V mucosu is quite aggressive and often grows over
crustose algae, such as Hildcnhruntia and Lithorhamiiia. It also is
commonly found growing over other Vermcanae where the penthe-
cia of the overgrown material project through the thallus of I' iimaisci
causing a potential source of misidentification.

Spores of V mucosa are not distinctive m either shape or size, San-
tesson (1939) reported the size of spores to be highly vanable. 7-15
X 4-8/im.

Pycnidia are quite common on V' iimcosci thalli. They appear as
slits ringed with brown and occur widely scattered or in clusters. It
is not uncommon to find mounds made up of aggregated pycnidia.

GENERAL DISTRIBUTION: Sweden. Norway. Finland, Ice-
land. Faroe Islands. Germany. Ireland. France (Santesson 1939).
England (Ferry and Sheard 1969). Wales (Fletcher 1973a). Green-
land. Sibena. Fuegia. Aukland Islands. Campbell Island. New
Zealand (Lamb 1953). Bntish Columbia (Brodo MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MASSA-
CHUSETTS: Es.se.\ Co.. Plymouth Co. NEW HAMPSHIRE:
Rockingham Co. MAINE: Cumberland Co. . Sagadahoc Co. . Han-
cock Co., Lincoln. Co. (MemI MSC). Washington Co. NOVA
SCOTIA: Colchester Co.. Cumberland Co.. Yarmouth Co.. Digby
Co.. Sheiburne Co., Halifax Co., Victona Co.. Cape Breton Co.
NEWFOUNDLAND: Northern Peninsula Section. East Coast
Section. Avalon Section.

Vcnucana .siruiiuUi is most noted for the strikingly large carbo-
naceous structures termed juga by Santesson ( 1939). The juga rise
above the thallus and tend to be broad and often branched. They
commonly encompass a perithecium. Near the edge of the thallus
they often radiate out from the center, marking the leading edge of
thallus lobes. They are different from the ridges of I' cnchsenii and
V. ditmarsica. being more like flattened plateaus and buttes than
narrow ridges.

The thallus is normally a bnght green when fresh but darkens in
more intense sunlight. It fades quickly in the shade. In the herbar-
ium the thallus soon becomes colorless and scarcely visible, a con-
dition causing early workers to think that the juga were the entire
thallus (Santes.son 1939). The thallus is continuous but often shows
necral cracks upon drying.

The penthecia are often amorphous, coalescing v\ith juga or
appeanng as spheres frequently with flattened tops. The perithecia
often are cracked or dissected and usually a shiny black.

In the shade inodification the perithecia become more nearly
spherical or may be elongated vertically. Juga become reduced,
causing the shade modification of V. sihatula to resemble V. ditmar-
sica. However, even in the shade modification, juga usually retain
their flatness and are reduced in size, but still are broad by compari-
son w ith other ridged species. Perithecium diameters are in the gen-
eral range of 0.05-0.3 mm. Zschucke ( 1934) listed the spore range
as 8-12 X 4-5;im and Enchsen ( 1957) as 8-1 1 x 4-5^m.

GENERAL DISTRIBUTION: Sweden. Norway. Denmark. Ice-
land, Ireland. France. Spain. Portugal (Santesson 1939). Wales
(Fletcher 1973a), Scotland (Brodo MSC). England (James MSC).

Verntcaria silicicola Fink in Hednck. Mycologia 25(4):305. 1933.

DESCRIPTION: Thallus brownish, continuous to patchy, not
areolate. Perithecia shallow domes to hemispherical. 0. 15-0.4 mm
broad; excipulum hyaline to dark below. Spores hyaline. 18-22 x
8-9,(111.

The author's knowledge of this species is limited to the examina-
tion of available herbanum specimens collected and known only
from Long Island. N.Y. The material seen resembled V microspora
to a considerable degree. The thallus was brownish and patchy. The
penthecia. for the most part, were shallow domes. Hednck (1933)
gave 0. 15-0.4 mm as the diameter of penthecia and those seen by
the author fell within this range. The salient charactenstic of this
species is the spore size, listed by Hedrick (1933) as 19-22 x 8-
9^m. The author was unable to find spores but Brodo (1968)
reported the range as 16-25 x 6-l0,(iii.

NORTHEASTERN AMERICAN DISTRIBUTION: NEW-
YORK: Suffolk Co. (Latham MSC), (Brodo MSC).

Vernicana siriatida Wahlenb. in Ach. Suppl. Meth. Lich. 23.
1803.

DESCRIPTION: Thallus entire, light to dark green usually
opaque w hen dry. more translucent when wet. containing numerous
black, rather broad ndges often branched, ridges especially com-
mon at thallus margins. Perithecia hemispherical to globular, often
irregular, flattened and/or dissected, shiny. 0.07-0.3 mm diameter,
excipulum hyaline to entirely dark below. Spores ovoid, may be
pointed at one end. 8-10 x 4-6fim.

NORTHEASTERN AMERICAN DISTRIBUTION: MASSA-
CHUSETTS: Essex Co.. Plymouth Co. RHODE ISLAND: New-
port Co. NEW HAMPSHIRE: Rockingham Co. MAINE:
Cumberiand Co.. Hancock Co. NOVA SCOTIA: Yarmouth Co..
Digby Co.. Sheiburne Co.. Halifax Co.. Victona Co. NEW-
FOUNDLAND: West Coast Section, Northern Peninsula Section.
East Coast Section. Avalon Section. NEW BRUNSWICK: Grand
Manan (W.R.Taylor MICH).

Genus Xanthoria

Xamhona candclanu (L.) Th. Fr. Gen. Heterolich. Europ. 61.
1861. Lichen candelahus L. Sp. PI. 1 141. 1753.

DESCRIPTION: Thallus foliose. light gold, diffuse or forming
rosettes, KOH + purple. Lobes small (0.2-0.5 mm broad) plane to
convex, lacerated, sorediate. soredia mostly apical, lobes tend to
nse off substrate. Apothecia ca. 2 mm broad with thalline margin.
Spores eight perascus. polanlocular. 9-13 x 4-6/(m.

This species supert'icially resembles members of the genus Cande-
laria for which it is named but differs in reaction to KOH and spore
type. The Candelariae are KOH- and the spores are not polanlocular
There is also a resemblance between Xanthoria candelaria and X.
fallax when the latter displays lobe widths at the narrow end of the
range for that species. However, in such a case the location of soredia
will distinguish them. In X. candelaria the soredia are apical or lami-
nal as opposed to the labrose mode in X. fallax.

GENERAL DISTRIBUTION: Alaska (Krog 1968). South
Dakota (Wetmore 1967). Nevada (Imshaug 1957). Washington
(Howard 1950). Utah (Wetmore MSC). California (Newcomb

21

MSC), Bnllsh Columbia (Ohlssim MSC). west central Canada
(Bird 1970). Cuba (Montague 1838-42). Greenland (Lynge 19.32).
Novaya Zemlya (Lynge 1932). France (Harmand 1905), Gerinany
(Erichsen 1957). Nepal (Awa.sthi 1965). England (Ferry and
Sheard 1969).

NORTHEASTERN AMERICAN DISTRIBUTION; MAINE:
Cumberland Co.. Sagadahoc Co.. Hancock Co. MASSACHU-
SETTS: (Fisher MSC). NEW YORK: (Brodo 1968). NOVA SCO-
TIA: Halifax Co.. Yarmouth Co.. Digby Co.. Shelburne Co.

NORTHEASTERN AMERICAN DISTRIBUTION: NOVA
SCOTIA: Cape Breton Co. NEWFOUNDLAND: Avalon Section.

SELECTED BIBLIOGRAPHY

Xaiitlwriu dedans [Lmk) Th. Fr Lich. Arct. 69. I860. Lichen ele-
.?(//;.< Link. Ann. Naturges. 1:37. 1791.

DESCRIPTION: Thallus foliose. red orange to citnne. nsing off
of sub.strate at least at tips. KOH + purple, forming ro.settes about
2-5 cm diameter, lobes nodular-conve.x. including tips. Apothecia
orange yellow to red orange, about 1 -2 mm broad. Spore colorless,
polarilocular. 10-13 x 5-8/(m.

Upon cursory inspection this species may appear to be crustose
but it does have a developed lower cortex and lifts off the sub.strate
at least at the tips. The lobes have a tubular appearance but of quite
irregular width and thickness, giving it a rather nodular appearance.
Like X. pcirietiiui it is devoid of soredia.

GENERAL DISTRIBUTION: Washington (Howard 1950).
Alaska (Krog 1968). South Dakota (Wetmore 1967). New Mexico
(Rudolph 1953). Anzona (Nash MSC). Colorado (Wetmore MSC).
Wyoming (Shushan MSC). Montana (Imshaug MSC). Iowa
(Imshaug MSC). Michigan (Imshaug MSC). Minnesota (Fink
MSC), Wisconsin (Malachowski MSC), New York (Brodo MSC),
Mexico (Beaman MSC). Manitoba (Gillis MSC). Northwest Tem-
tory (Presi MSC). Nova Scotia (Lamb 1953). Greenland (Lynge
1932). Finland (Rasanen 1927). India (Awasthi 1965). China
(Magnusson 1940), New Zealand (Nylander 1888). Ontario (Wang
MSC), Iceland (Copeland MSC).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE:
Sagadahoc Co., Hancock Co. NOVA SCOTIA: Digby Co.,
Shelburne Co., Halifax Co., Victona Co., Cape Breton Co. NEW-
FOUNDLAND: West Coast Section, Northern Peninsula Section.

Xanthoiui pancniui (L.) Th. Fr Lich. Arct. 67. 1860. Lichen
panrimi.s L. Sp. PI. 1 143. 1753.

DESCRIPTION: Thallus foliose. orange to citrine, radiate,
often rosettclike. KOH + purple, lobes flat, wnnkled. ca. 1 mm
broad, usually forked at the tips, soredia absent. Apothecia numer-
ous, ca. 5 mm broad, plane to concave with thin thalline margin
disappearing with age. Spores eight perascus. polanlocular color-
less, 10-13 X 6-9,(01.

This widespread oceanic species Is easily distinguished from all
other littoral Xanthoriae by its relatively broad, thin, distinctly
wrinkled thallus divided dichotomously at the tips. The apothecia
often contrast with the thallus by being more red colored,

GENERAL DISTRIBUTION: Greenland (Lynge 1937). Nov-
aya Zemlya (Lynge 192S). Sweden (Dcgelius 1935). Finland
(Rasanen 1927). Italy (Jatta 1909-1911). India (Awasthi 1965).
Cuba (Montagne 1838-42). Tenenfe (Imshaug MSC). England
(Ferry and Sheard 1969). Wales (Fletcher 1937b). Spain (W. & C.
Culberson MSC).

AWASTHI. D D

1965. C;ikili>guc ol the lichens Inmi India. Nepal. Pakistan, and Ccvlon
Nova Heduieij. Bcihcftc. SuppI 17. LIT p
BIRD. C D

1970 Keys to Ihc lichens ol wesi Lenlral Canada The Hcrhanuni, Dep
Biol . Univ, Calgary, mimeogr . p 132
BRODO. I M

,1968. The lichens ot Long Island, New York A \egelaIional and llonstic
analysis NY Stale Mus .Sci Serv Bull 411). .V'Op
DKGF.LIUS. G

19,'l.S Das o7.-anische element der strauch-und-lauh-tleehlen flora von Skan-

dinavien Acta Phytogeogr Suec 7 1-411
1939. Die tiechlen \on norra skatton Lippsala L:niv .Arsskr II I-2U.S
ERICHSEN. C F E

1957 Flechten Mora von Nordwestdeutschland Gusta^ Fischer Verlag.
Slultgan. p 1-41 I
FERRY. B, W,. and J. W SHEARD

1969, Zonation of supralitloral lichens on shores around Ihc Dale Peninsula.
Pembrokeshire Field Siud 3 41-67
FLETCHER. A

1973a, The ecology of marine (littoral! lichens on some rocky shores of

Anglesey. Lichenologist 5:368-400
|473b. The ecology of maritime (supralitloral) lichens on some rocky shores
of Anglesey, Lichenologisi 5:401-422
HALE. M E . Jr . and W L CULBERSON

1970 A lourth checklist of the lichens of the continental tinned States and
Canada Bryologist 73:499-543.
HARMAND. L J

1905, Lichens de France Epinal. Pans 4 609
HARRIS. R

1975, A ta,vonomic revision of the genus Arthopyrania Massal S, Lat (Asco-
niycetes) in Norlh America Mich Slate Univ . East Lansing. 301 p
HEDRICK. J

1933, New genera and species of lichens from the herbarium i.A Brxice Fink
I, Mycologia25 303-316
HOWARD. G E

1950 Lichensol the slateol Washington Lmv Wash Press, Seattle, 19 I p
IMSHAUG. H A

1957, Alpine lichens of western United Slates and adjacent Canada I The
macrolichens Bryologist 60: 177-272,
J,«iTTA. A

1909-191 I Flora italica iryplogjma Pars III Lichens Rocca S Cus-
ciano Stabilimento Tipografico L Cappelli, 460 p
KROG, H

1468 The macrolichens ol Alaska Nor Poljnnst Skrl44, ISOp
LAMB. I M

1953 Antarctic pyrenocarp lichens Discoverv Rep 25 I 30

1954 Lichens of Cape Bretim Island. Nova Scolia ,^nnu Rep Nail Mus
Can 132 239-313

L'l NGE. B

1928 Lichens Irom Novaya 7eml>a level ol A: cinnpi'iii and
Uiaiwnn Rep Set Results Norw Fvped N. naya /cmKa 192 1 , 43, 294

P
1432 Lichens Irom norlh cast Greenland, II .Microlichens Skr Svalbard

Ishavct SI 1-143
1937 Lichens Irom west Greenland, collected chielly by Th M

Fries Medd GiOnl I IHlSI 22s ,,
MAGNUSSON. A H

1932 Lichens Irom western Northamenc.i, mainl> Washington and

Alaska Ann Cryptog Hxot 5lll:l6-38
1440 Lichens Irom cental Asia Rep Sci hvped Norlh w Prov China 13

I6S p
MONTAGNE, ,1 F C

1838-1842 Botanique-Plantes Ccllulairies In R de la Sagra, Hlstoire phy -

siqiie politique et n.ilurelle de I'lle de Cuba 8 105-237,

22

NORDIN. I

1972 Ciilop/atii ^i'Ji Gaspurnnij 1 Nordcurupa Upps.il.i.p 1 1K4.
NYLANDER, W.

1888. Lichencs Novae Zelandiae Pans, p l-15h
I8W Lichencs Japiiniac Pans, p l-i:2.
POELT, J

1959. Bcstimmungsschlussel Europaischer Flechien. J. Cramer. Lehre, p
1-259
RASANEN, V

1927 Uber Flethtenstandorte und Ficchlcnvegelaliiin im wesllichcn Nord-
finland Ann Soc Zool Bui Fenn Vananui 7 1-202.
RUDOLPH, E D

\9^} A contribution to the hehen flora ol An/ona and New Mexict) Ann
Mo Bol Card 40:6.^-72
SANTESSON, R

1939. Amphibious pyrenolithens I Ark Bol 29AI 1(1) 1 67
SERVIT. M

1954. Ceskoslovenske lisenjnilsy celed: Verrucariaceae Nakludalelstvi

Ceskoslovenske Akademie Ved Prana

SWINSCOW, T D V

1965. The marine species ot Atlht>[)\rt/iui in the British isles Lichenologist
3 55-64.
TUCKERMAN. E.

1882 A synopsis ol the North American lichens: Parti., comprising the Por-
nieliiui'i. CkititmH-i, and Covn(ii;tinifi ,S E Cassino. Boston. 261 p.
WADE. A E

1965 The genus G//<v'/'"" Th Fr in the British isles Lichenologist 3 1 -
28
WEBER. W A

1962 Environmental modification and the taxonomy of the crusto.se
lichens. Sven Bol Tidskr. 56:293-333
WETMORE.C M

1957 Lichens ol the Black Hills of South Dakota and Wyoming Publ Mus.
Mich. State Univ . Biol Scr 3:209-454
ZSCHACKE. H

1925 Die mitteieuropaischen Verrucariaceen IV. Hedwigia 65:46-54
1934 Epigloeaceae, Verrucariaceae und Dermatocarpaceae Rabenhorsl's
kryptogamen-llora 1 . part 9, 695 p

23

SYSTEMATIC INDEX

Anhopxrciua hulodyws 1.2.3.9. 15

Calopkuci marina 8 . 15

Caloplaca verruculifeni 8. 16

Caloplaca microtluillina 8. 15. 16

Caloplaca scopulari.s 8. 16

Lecanora f>ranlii 2.1 . 16

Lichiiia amfinis 2.6

Stigmidiiiin marinum 9. 16

Verrucaria amphibia 12. 13. 17

Vernuaria ceulhocarpa 10. 17. 18. 19

Verrucaria degelii 3. 12, 13, 18

lirrucaria ililiiiarsica 2.3. 14. 1 8. 20. 2 1

Verrucaria erichsenii 2.3. 13. 14. 18. 19.21

Verrucaria iiUeniii>rescens 10. 17. 19

Vernuaria maiira 3. 10. 12. 15. 16. 17. 19. 20

Verrucaria microspora 3. 10. 17. 20. 21

Verrucaria nnaosa 2.4. 10. 17. 20. 2 1

Verrucaria silicicola 10.17.21

Verrucaria srriatula 2.4. 15, 19. 20. 2 1

Xanihoria caiulelaria 6.21

Xauthoria elegans 7. 22

Xanihoria parielina 7, 22

24

ACKNOWLEDGMENTS

Preparation of the "Manne Flora and Fauna ol the Northeastern United
States" is being eoordinated by the following Board:

Coordinating Editor; Melbourne R. Carriker, College of Marine Studies,

University of Delaware, Lewes. DE 19958.
Editorial Advisers: Marie B. Abbott. 259 High Street. Coventry. Conn.

(Formeriy with Gray Museum. Manne Biologi-
cal Labtiratory. Woods Hole. Mass.)

Arthur G. Humes, Boston University Marine Pro-
gram, Marine Biological Laboratory, Woods
Hole, Mass.

Wesley N. Tiffney, Retired, Boston University, 226
Edge Hill Road, Shanan. Mass.

Ruth D. Tbrner, Museum of Comparative Zoology,
Harvard University. Cambridge, Mass.

Roland L Wigley, Retired, National Manne Fish-
enes Service. Northeast Fisheries Center.
NOAA. Woods Hole, Mass.

Robert T Wilce, Department of Botany, Univer-
sity of Massachusetts. Amherst, Mass.

The Board established the format for the "Manne Flora and Fauna of
the Northeastern United Stales," invites systematists to collaborate in the
preparation of manuals, reviews manuscnpts, and advises the Scientific
Editor of the National Manne Fishenes Service.

Thanks are expressed to Henry A, Imshaug for his assistance in
research and advice in prepanng the manuscript. Thanks are also expressed
to Melbourne R. Camker for continued assistance in later research stages
and manuscript preparation. Richard D. Wood was very helpful in the field
work in the Narragansett Bay area. Helen S. Taylor, Brian M. Taylor, and
Sheryl M. Taylor contnbuled both in the research for and preparation of the
mauscnpt. Adele Denison and Ralph Gorton aided in the preparation of the
final manuscnpt.

25

COORDINATING EDITOR'S COMMENTS

Publication of the "Maiine Flora and Fauna of the Northeastern United
Stales" is most timely in view of the growing univeral emphasis on environ-
mental work and the urgent need for more precise and complete identifica-
tion of coastal organisms than has been available. It is mandatory, wherever
possible, that organisms be identified accurately to species Accurate scien-
tific names unlock the great quantities of biological information stored in
libraries, obviate duplication of research already done, and often make pos-
sible predlctionof attributes of organisms ihat have been inadequately stud-
ied.

Ronald Taylor started his research on lichens in l%4 at Michigan State
University where he later received his doctoral degree. He was then, and is
now, on the faculty of Lansing Community College at Lansing, Mich.

Preparation of this manual was supported in part by a grant from the
Environmental Protection Agency to the Edilonal Board of the "Marine
Flora and Fauna of the Northeastern United States." Work on the "Manne
Flora and Fauna of the Northeastern United Stales" by the Coordinating

Editor IS supported by the College of Manne Studies. University ot Dela
ware.

Manuals are available from the following:

Supenntendent of Documents. U.S. Government Pnnting OITice, Washing-
ton. DC ;a;02. for a charge.

User Service Branch. Library and Information Sen. ices. DiMsion D822.
Washington Science Center. Building 4. Rockville. MD 20852. at no
charge as long as the supply la.sts.

National Technical Information Services. U.S. Department of Commerce.
5285 Port Royal Road. Spnngfield. VA 22161. cither as paper copy or
micronche. for a charge.

Manuals arc not copynghted. and so may be photocopied from the NOAA
Technical Report NMFS Circulars available in most major libranes.

The manuals so far published in the NOAA Technical Report NMFS Cir-
cular series are listed below by author, title, circular number, and NTIS
accession number.

Manne Flora and Fauna of the Northeastern Linited Slates:

COOK. DAVID G, and RALPH O BRINKHURST Annelida: Oligochaeta

BORROR, ARTHUR C Protozoa: Ciliophora

MOUL, EDWIN T Higher Plants of the Manne Fnnge.

McCLOSKEY, LAWRENCE R Pycnogonida.

MANNING, RAYMOND B. Cru.slacea: Stomatopoda

WILLIAMS, AUSTON B. Cr\istacea; Decapoda

POLLOCK, LELAND W Tardigrada.

LARSON, RONALD J. Cnidana: Scypho/oa.

CAVALIERE, A. R. Higher Fungi: Asconiyceles, Deuteromycetes, and Basidiomycetes.

COULL, BRUCE C. Copepoda: Harpacticoida

CUTLER, EDWARD B Sipuncula

PAWSON, DAVID L. Echinodermata: Holothuroidea.

HO, JU-SHEY. Copepoda: Lernaeopodidae and Sphynidae.

HO, JU-SHEY. Copepoda: Cyclopoids Parasitic on Fishes.

CRESSEY, ROGER F Crustacea: Branchiura

BOVEE, EUGENE C, and THOMAS K SAWDER Protozoa: Sarcodina: Amoebae.

WATLING, LES. Crustacea: Cumacea

ZULLO, VICTOR A. Arthropoda: Cin-ipedia

CAIRNS, STEPHEN D. Cnidana: Scleraclinia.

TODD, RUTH, and DORIS LOW. Protozoa: Sarcodina Bcnlhic Foraminifera

BUSH, LOUISE F Turbellana: Acoela and Nemertoderniatida.

TAYLOR, RONALD M. Lichens ( Ascomycetes) of the Intertidal Region.

Circular

No

374
378
384
386
387
389
394
397
398
399
403
405
406
409
413
419
423
425
438
439
440

NTIS NO.

COM 73 50670
COM 73 50888
COM 74 50019
COM 74 .50014
COM 74 50487
COM 74 51 194
PB 257 987
PB26I 839
PB 268 036
PB 268 714
PB 273 062
PB 274 999
PB 280 040
PB281 969
PB 222 923
PB 285 538
PB 296 460
PB 297 676
PB 124 520
PB 225 053
PB 219 387

26

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