NOAA 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

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 MF Circular I, part ur tht: sub'>t:nc, "Munne l'lora ,mu hlunil 01 the Northe.t\tt:fn Unlteu States." ""hlch t:onSI,\', of onglnal tlillstruteu 1llout:rn Illunuuls Oil thl' Idcntlfll;atloll. ,Ia .. 1Il<.:.ltll1n. ,mu general bIology 01 the e,tuanlle anu (oustal 111anl1l: plant' olnu anl111ul, of th~ northeastern lJnlh!O States The 111anuab arc puhh,heo at Irregular Intcnal, on 01' man) tJ\a of the regIon a, therc are peel.lil,t" u\ atl able to collaborate tn theIr preparatton

GeographIc coveruge of tht: "Manne Flora oIllli Fauna pf thl' ortheu,tt:rn l 'nlteu St.lte .. t\ pl.tnn d hl 1I1clude organl'Il)!, fro111 the heau""aters 01 e,tuClne, ,eav. ,mi til appro\llllutclj the 2()() III ucpth (In the on· ttnental shelf from MaIne tll VlrglnlJ. but ma) \.If) '''111ev. hat v. Ith e.lch rnaJIH t.1 nn ,lI1U thc Interest (If collaborators. Whene\ er po"lblc rcpre,cntatt\c 'pelI111ell' uealt v,lth III the manual arc ucp<.' ItCU In th reference collecttons of major museum, 01 the rcglon

The "Manne Flom anu Fuuna 01 the nrtheastcrn Llnlleu Stoltes" I Oclng prep.1red In ,oll,lh.,rallon "'Ith S) stematle SpeCiultsh In the l;l1Iteu tatc, and abr'lau tach rn,tnualls t1,l';cd pnrnanl) (In re ent and ongoing rev ISlonal) ') ,ternattl re,carrh and a ire,h nJmlnatm pf the pl..tnts and anl111,11 E:l h 111,lJor ta\on. treated In a separ..ttc manual. Incluoe, .In Introducttnn. IlIu,trolteu gil" ,II) unll 'rill unglnall) tllu trated keys. annotated ,hecklt,t v. Ilh inll'rrnatl\lO ,\ hen ~\ atlahk llll uI,tnhutIPn. hablt.lt. Itfe hi tol) nd related blOlog). references t~) the major IttCI1lturc "I the: group. and ~ ') -telllati Ind \

These manua" are intended Illr u,e h) blolog) ,tudent,. blllil'gl'h. bl(,logleal oceanographer.;. 1I1formed la)men. ano others v. I,hlng to lu<.'ntll) C".I,lal nrg .. nl,m, ror thl' reg Inn Ott nth) ,.In \Cne gUides to addttl~)nal 1I1f lrmatton a!xlut 'pee It.!' pr gmup,

The manual> arc an outgrov. th llf thl· v. Id<.'l) u,cd .. K<), t" \loInn< Jill <.'rtcbr .. tcs 01 the \\Oods Hole RegIon." edlled b) R I mllh 111 1\)04. and pnlduud under the oIU pIce 01 the S) tem,llI holog) Pro· gram. Manne BIologIcal L.1bt.,ratof), Wlxlth Hllk. f,la ,\It<.'r a ulll lent number 01 manu.l! 01 related taxonomIc groups have been publl\hed. the manual, "til be ft.!' I l'd. gr upt'd. nd h u<.'d pe~1.l1 \01· umes. \\ hieh v. til con'l,t ll! ':ompliatllln, I~lr ph fa or gn>up, llf ph) I

CONTENTS

Introduction ..... .. ...... . .. . .... . .... . . . . . ................... .. ..... . .... . .. . ...... .. ................. . . Methods of collection and studying . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 1 Ecology ......... .. ..... . . ' ........... . .. . ..... .. ....................... . .. . . . ......................... . . 2 Glossary . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . 4 llIustrated key to species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Annotated list of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 15 Selected bibliography . ...... . ........ . ...... . ........ . ..... . . . ....... . .................................... 22 Systematic index .... . .. . ............. . . .. ........ .... ......... . .............. . ........ . .................. 24 Acknowledgments ...................... . ................... . .......................... . .................. 25 Coordinating Editor's comments ........ . ................................................................ . ... 26

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

III

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

the Intertidal Region

RONALD M. TAYLOR I

ABSTRACT

This manual treats the lichens found in the intertidal region from New Jersey to Newfoundland. Methods of

collection, preparation. and study are briefly treated. Twenty-two species are covered, both in an illustrated key

and an alphabetical li sting, with brief descriptions and notes on ecology and distribution. A glossary of terms is included .

INTRODUCTION

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 lit­erature and from inquiries directed to the author that the marine 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 or growth form com­prised 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 lhe marine lichens treated in this manual, except for the genus Lichina , are fungus dominated and the fungus is an Ascomycete.

Marine I ichens occur in three growth forms: fruticose, foliose, and crustose. Fruticose forms are usually upright or pendant in rela­tionship to the substrate; they stand out away from the substrate. LichinQ, 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 Xanrhoria . 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, Arrhopyrenia halodyres , 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 facultative lichen is its occurrence with, or at the same height on the shore as, an obligate marine species.

IScie nce Department, Lan>lng Community College. LanSing. Ml 48914.

METHODS OF COLLECTION AND STUDYING

Marine 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. Unfortunately, the specimens are often wet, and the wet bag may need to be placed in another bag. PIa tic 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 transfer­red 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 causes excessive reticulation that is detrimental to fu1!!re stu<ry.

Hydraulic devices are available which break rocks by applying pressure to chisel like 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 frag­ments 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. Bright illumination is important because of the dark color of many thalli, especially in the genus Verrllcaria. Thalli should be observed in both wet and dry conditions. Water added to the thallus sparingly to avoid unnatural reticulation often increases its trans­parency, thus revealing subsurface patterns.

Secltnn, 01' Il11illng \II1.IClUrl'\ 111,1) Iw 1lI.llk h) h,llld II I1h rJlnf bladc rhc sl11all \I/C ,)1 I Ill' pellll1l'll.l "I 111,111) Pl'll~~ 111 I, I

rl/('(//"/o an: dilTinill hUI [hI( IlllP'''\lbk In 'L'lll<)[1 ,\ d"uhle CII· d rJlOf hladc, hn)l..L'n kl1glhlll\l', 1\ 111<1\1 1111 .. hll'

l\1ll'n)I{)ml' 'l'l'liOI1' ,lfl'l'omplll' ,l1l'd h) Ihe 111IIlllll III e n,llure of the Ihalli and 1IlL'1I ,ldhl'll'llL'l'l<> the 10 ' 1.. IIh Ifltc Illrth ITIlore the inH\lll,'rell,1 (11 PYll'lhk',If11<lU' II, hl'l1~ .11'1.' h.tr I .II1J bnl1le md dOIH11 rcspnnd 11,'1\ I" Ihl'll\llal p,lrtlllllllllhc Idll1 t~dllllqUl'

1\1{1 tl','hnlljlh,'\ l'mph,) 111£ 1 In:all1g 1111 r\l(()I11~ rc .11I~f.1

lOr, Oil" technlljuL' II110h " !,(lllnn '<I 11.1110\\ 1.1) er of .1 '.If 11\

smalll'{)lllalnCI. rhl' pcntllL'lllllll IS 11101 1(11 d C\l'mlllll1l' "Ilh

8(1lh t~,hm'lu( ha\l' Ihe dl oId\oIntol dues lIn the ~hde urroundmg the c lion the slide i. to be photogr..lphed

"1\1. 1111 d 11 ohol

n Ih Ind

If

ome SPCCIC uf hchen, gl\ a (olor reaction I pow lum h)dr xide, A drop of 10% olullon I .Ipplt d II Ith 01 dmppm pipelle or capillal") tubc, dlre..!l) to th c nc! f to rn dullJ exposed b) fmgmentoltlon or an tn(1 Ion

ante son (1439) dc, nbed a number 01 em lronm ntall mduced morphlllnglcal \ an,llIun tn Itttural hch n In 'en ntl .md described m delall the \ analllln, LlI c nmn c!eted pelle B th Weber (1962) and Sante. on (II) 9) hJ\ L'mph.1 \Zed Ihe Impor lance of undeNand IIlg the rang of \ anJlIon tndUl d b) Ih em ironment.

Thc author ha. tml'ed \ anaUllil of mglc thallt n m (lntlnU­ous to areolate and lrom smooth tll ndged rhL' auth"r 11.1' .11 0

obseryed I'ariatlons 01 perithecla frnlll ,unken I" pnlllllnL'ntl) raised on a ~ingle thallu., E\en [he darkne {,fthe I!l\\er Ide ,)fth' excipulum (Illner llIllllg of pentheClulIl) I. yU([L' \ ..lriable \\ uhlll a single speCIes. Shade modlfll:atlon I \\ ell I..nn\\ nand predlet,lbk among littoral lichens Shade eauses Ilghler plgmcnt.ltllln and redUC[lOn in size and number of Juga (blad pegs, lump', ()[ ridge protruding from the surface) Some 'pecles lhat are di\linct in appearance when growing In good 11Iuminalion app.:ar much Ie distinct when growlIlg III the shadc, The dIstinguishing fealure 01 a species are frequently reduced by the ,hade modification , As a result there is considerable taxonomIc confusion among the hade modifications of Verrucaria sTriatu/a. r: dillllllnico, and. to :1

lesser degree, V erichsenii, Too much ~[res~ can be placed on perithecium size. Vanatlon of pentheclUm size on a ~lIlgle thallus beyond the range accepted in some taxonomIc I..eys have been observed, Thus the taxonomic character "perithecta IIghll) smaller (or larger) than . , . ,. should be regarded II Ith u~piclOn ,

dl pc I

h..lhll I , II I n I

lOlilu tn· \.m tlon III Ih

'pccle

nh rn end of th luJ~

mn~ • \pl.ltn

til c dteL!' \\ L're compu[L'd <Ire altnll), air Jnd \I .I[d I mpc r.lIure • ,,)IM IIlS11Iatl,'n, and ud.11 r.lIlge ~l' Impie In II ~ rs em~ rged c'uhd 1M the suc,'c" {If .In) P<:CIC' l'r I'M gencr.ll dl\ ersll) H,)II el cr. b) tall) ing [hL' number {,r lime, Ihal e..lch l:nl tnlnmental laCI,lr II a I III ol\l:d as one "I [he mo. [ Imp 'T1ant mtluence,. Ihe f{lllol\ Ing fre­quen'::lc re,ulted: ,allnll), 21, In,,,la[llln. I:, aIr temperalure, 12, udJI range, 7. and II aler Ie I11P'::ralU re, 2,

Wher.:: suffiCient collec:u{ln, \1 ere made II) pnn idL' adequale data, \erllcal dbtnbullons 01 Ihe lanou, marine lichens I\ere graphed (Figs. 1- ) In Ihe,c gmphs the eXlent abo\ e mean high

50~

50:

50~

1()n

'5m

MHW

4hrs

Shrs

12hrs

16hrs

20hrs

24hrs

Figure I.-Vertical di stribution of Arthopyrenio ha/odytes.

10m

5m

MH '..J

4hrs

Shrs

12hrs

16hrs

20hrs

24hrs

Figure 2.-Vertical distribution of Verrucaria degelii.

l()n

5m

\~H(.,!

4hrs

Shrs

;---12hrs

16hrs

~ 20hrs I 24hrs

Figure 3,-Yertical distribution of \ 'errucaria ditmarsica.

50" 50:

50~ 50~

50 50'

......... "-..

"-

" . t.hrs

hrs

12hr'

16hr s

28hrs

24hrs \

Figure 4. - Vertical distribution of Verrucaria enchsenil.

l()n

5m

MHW

4hrs

Shr

12er

16~ rs

Figure S. - Vertical distribution of l errucaria maura .

1

I t .. rs

Figure 6. - Yertical di,tributi on of \"rurana mIcro para

50:

l 10m

5m

MHW

4hrs

8hrs

12hrs

16hrs

20hrs

24hrs

Figure 7.-Vertical distribution of Verrucaria mucosa.

tide is given in meters . Due to the great variation in the distance between mean low tide and mean high tide (ca . 1 to 10 m) , the dis­tance below high tide is given in hours of submergence. Each inter­val on the X-axis represents 10 % of the total collections of that species by the author. In each figure the graph on the right repre­sents the vertical distribution 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.

laminal soredia~~~~

upper cortex lower cortex

lam

5m

50% 50

~l HW

4hrs

8hrS

12hrs

16hrs

20hrs

& 24hrs

Figure 8.-Vertical di stribution of Verrucaria striatufa .

Apical At the tip or apex (Fig. 9). Apothecium (pI. Apothecia) Disc, saucer, or cup shaped fruit­

ing body (Fig.9). Areolate Broken by clefts into small , usually angu lar and irreg­

ular patches (areoles). Areole (pI. Areoles) Area of a thallus divided from the rest of

the thallus by fissures (Fig. 10) . Arete (pl . Aretes) Sharp peak elevated above a ridge .

Ascocarp (pI. Ascocarps) An Ascomycete fruiting body giving rise to and containi ng asc i and ascospores (Figs . 9 , 10).

Ascus (pI. Asci) A sack-like structure within which ascospores are formed and contained in the ascocarp (Fig. 10).

Carbonaceous Made up largely of carbon deposits .

soredia

.'" ,".

'. 'f ! .. _

thall ine margin

Figure 9.-Composite foliose li chen with apothecia.

Citrine Lemon yellow color. Conidium (pI. Conidia) Asexual reproductive spores of Asco­

mycete . Continuous Thallus unbroken by gaps or fissures . Cortex (pI. Cortices or Cortexes) The compact outermost fun ­

gallayerofa lichen thallus (Figs. 9,10). Craterous Having a depression resembling the crater of a vol­

cano. Crenulate 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) .

Cusp (pI. Cusps) Crescent shaped mound . Dendritic Having a pattern resembling the branching of a tree

limb . Dispersed Scattered . Endolithic Growing within the rock substrate . Epithecium (pI. Epithecia) Covering of the hymenium. Evaginate To protrude as if a pocket turned inside-out. Excipulum (pI. Excipula) As used here, the inner wall of a

perithec ium . Foliose A type of thallus , leaflike or lobed , often with rhizines

or hairlike processes on lower surface, usuall y with lower cortex , algal layer limited to upper side (Fig . 9) .

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

FUSiform Narrowed both ways from a swoll en 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 (pl . Hymenia) The layer of the ascocarp contai ning

the asci and usually paraphyses (Fig. 9). Hyporhal/us (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. In volucrellum (pI. In volucrella ) The usually bl ack and carbo-

areole~~~ ____ ~_

naceous covering of a perithecium (Fig . 10) . Isidium (pI. Isidia ) Small cylindrical, coralloid , or globose

outgrowths of the thallus serving as vegetative disseminules . They are covered with cortex and contain both fungal and algal compo­nents (Fig. 9).

Isthmus (pl. Isrhmuses) A narrow passage. Jugum (pI. Juga) Protrusions of carbonaceous material extend­

ing at least partly above the thallus surface as ridges , pegs, or irreg­ular lumps (Fig . 10).

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

Lenticular An area, pointed at each end, inclosed by two inter­secting convex curves.

Littoral lichen Growing between the tides . The term is extended herein to include those struck by sal twater due to wave action and is used interchangeably with the term "marine."

Locule (pl . Londes) 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).

Morphotype (pI. Morphorypes) A thallus of a given species hav ing 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 (pl . 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. Parasymbionts) An organism living In

harmless association with another without mutual benefit. Pendant Dangling . Perithecoid In the shape of a perithecium .

Figure IO .-Composite crustose li chen with perithecia .

Perilhecilllll (pI. Perilhecill) A na\h.\hapeJ "'Ll)L'arp enLlmeu except for the apica l pore (ostio le), It 1\ u, uall y 1111)le or k,\ ,ul1h.el1 into the thall us (Fig, 10)

Phycohio/ll (pI. Phl'colnOIlI,\) The algal 111\':111her 01 the itLhell ymbiosis (Figs, 9, 10)

Pselldosepllllll (pI. PI(,/Illo,\('{llci) Appeanng to he .In IIlIl·1I1.d wall but Oot composeJ of the ~allle maten,d fnrnllng the LL'II 11;t11

P"cllidilllll (pi P"clllclill) mall f1a\h. \haped til glohlll.11 'tnll" ture resembling perithecI.\ but contal11l11g lll11iuI<l il1\te,lu 01 ,I'L I and ascospores,

PyrellcOCarlJOlIs Ha\J11g penthena . Pyrello/ichclI (pi P,r('//olich,'I1.I) I Ic hen h,l\ l11g pc'nthe l.t

(Fig, 10), Relllforlll Kldne) \hapeu. ReI/CII/alIOII A netllh.e appearance. Rilllose Haling minute Lr,lch.-. usuall) rcklTlng 10.1 lor111 III

areolatlon SO-HCO/OIIS ro\\lng on a roeh. or roe"·II"c \lIbstfJtL'. Sessile 01 supported b) a \tal h. SlIllp/e DesenblOg a spore h.11 109 a \Ingk, undl\ IdLd L,II It)

.\'n/n/lCllc 11 ,,\ Ill' SIH dl.l 1111 Ihc Ih.dlu~ SU/{,tll/ l l/1 (pi \oln/tll) mc,w, II( VL' 'ctatl\C rcpIllduUI Ifl "I

tl1\: I I dll' 11 LIH1\I,tln,' 01 111.I'SL'S 01 hyph.tc .mll Ihelr.1 SI l.tted ul ',1'

nlptulll1g Ihrough .Ind nol llllcrcd h} III !.onl,lImn' orlCX 1 Jotg (1)

,\'/1I1/1 :11/1/' 11'1 'nlll'\) J 11.11 MC,I .tho\c high IIUC lit II. 'Hrud b) drop' 01 C.tI\.lIer re,uliing Irol11 Ihc Imp.1 I I \\dICS 00 Ihe rOL") ,hllre IIndei 111111 !CHill Lllndtlll,n,

,\1(11111 IIJ/tl\ '(1111 I pi 'II/II II ( hJI ,HC,I, ,J IC Ihe u UJI pIa) lonc Ih.lt I Spl,l)cd \\ IIh , .I .... dler (,nl) dunng.1 torm

\11/" /1/111'\ H,lIrl) ,ur\cJ j'UIV. If 1 \111>,11111(' I { 1\ 1/1' h Hlnd.tne th.lt ar poorly II IIngul hed .\1I1>1II( IXI'd Sunken Cl111r I} or rno tl} bel \\ Ihe uppcr uria e

III Ihe th.tllu \111111/0111//(01 III H,I\ 10' ~oo In'110n at IOlenal 1\ 10 In

,lPPl.lf III I: n ,HI} like a Inn' of b\:ad but not \0 pmnouo~ed

/7/(/1/1111 111<1' I:/1I Ipl IJ/or '/11 ) ,\ pot he, tal nm hal 10 the am ('101'0 ilIOn ,\ the Ihallu (II I)

/11111/11 \ t pi l71a/l1) I hc It hen hod} ont,lImn !loth Ig.1l Joll lun '.tl omronent

/ilhcrC/I/(/I1 lId\ln' d "an} appear.tn e

ll.ll ~lR \ll . J)KL ' 10 PI ' 11 '-;

The ke) IS based pnmanl) ,10 L'har.tcten IlL' IhJt opc at 20 JTl;I mil allOn til me observatIOn of spores 1\ nh a c()mpnund nllefl)\l\)r~ 111~1) be requlr d

The \'ertl~JI sections of pcnthecia 1)1' I (IrtIC(1I /{/ ,pc I .\lu tmlLd 10 Ih "'\ rc I pi al lion Because of the e:\trcme degree or I anation III pelll' of mann' IlI.h n (e pcLlJII) of I am ana, Idenull alloo of pc Ie from Ih )

alone hould be regarded as (entallle unIt! the more d uti d de nptilln In Ih ,\nnut.1ted I I t of pc IC 1 read Five pecies from the Canadian 'lariume Pmllll<ee ,Ire til Iud d \ en thou h at pre ent the) rc n t nil. 0 frum th a tcrn Cnllcd

States, At least one species. lu-rucaria 1I1<1l1r,/ ha' nnl been Lollcued 111 Ih" ea tern L nil d SI.Ih: re entl) but the author hOI cn pc 1m n collected from the area o\er 50 ) r ago

2 (1)

Thallus fohosc (Fig. 9): a1\\a) orJngc "nd KOH ... pUI11I"

Thallus crusto,e (Fig 10) onl) In a lc\\ pC(,le l)ral1ge ,Ind KOH purplc

Thallu erect. fruticose. compo\ed pnmanl) 111 alpl (Ca[Vlltrll) filament, U 0 drp )1cnthnlOld Ihg II)

Figure t) ,-I.lch,no ranfinll, ~tale j, 0,1

mm.

Thallus with oredla rna tl) apical (Fig. 12). thallus lobe, rlOel) dn Ided (0 2-U. : 111m \\ Ide)

Figure l2.-Thallus or XO ll lh oria cOlldelorio, Scale i, t mm .

. .. U,/lIlIa ,ofljim

.\ (/I/I/,orio c(/Ildc/arta

2 (1) Thallus not sorediate, lobes wider than 0.5 mm . , , . ' .. , , ' , , . , .. , , , , , , , . , , , , ' , , . , . , , , , , .. ' , , , , . , . , , .. , ..... , . , . , ' ,3

3 (2) Lobes flattened and 1 mm or more broad (Fig. 13) . .. .......... . ................. .. ...... .. ... .. . . Xanthoria parietina

Figure 13.-Thallus of Xalllhoria parielilla. Scale is 1 mm .

3 (2) Lobes nodu lar-convex. lobes less than 1 mm wide (Fig . 14) .. . .... .. ..... ... ... .. ...... .. . . ... ..... . . Xanthoria elegans

Figure 14.-Thallus of Xalllhoria elegalls. Scale is 1 mm.

4 (1) Ascocarp an apothecium (Fig . 9); thallus light (shades of white , gray, or yellow orange) .. .. ..... . ....... . ........... . ... 5

4 (1) Ascocarp a perithecium (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 . 15),11-18 x 6-8.5J.tm .... . ....... . Lecanora grantii

Figure 15 .-0ne-ce ll ed spore. Scale is tOllm. C ~

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

Figure 16.-Polarilocular spore. Scale is IOl'nl.

6 (5)

6 (5)

7 (6)

7 (6)

8 (7)

Thallu covcred with globl)~c I\llila (Fig. 171. ~r(}rc\ C) 14 4 hi/ill

Fi~url) 17.-1 hiJllu, of (a/o"llInl If"'" "II/'-fU Slill,' " III Oil".

Thallu~ \\ Ithllut 1~ldia .. . ..

~jj!urc 1 .-1 h.lllu, nf (a/opl"," ,opul", ... ~t:.J1t' i ... 10 mill .

Thallus IfTcgular. comp'h~J ,'I ca!lcr~d 10 or nun ('r u granule

Thallu~ cltnnc of gmnule\ I-J 111m Jlan! I r (f I I I, pore I -I

Figure 19.-Thallu, uf Caloplaca m.ao.hal/llla. ~tal. j, 10 mOl

Ul"p[lII11 \i'lf IIUI I I r{/

7

11m ( alop/aca fcoplihm

5- I'm al pia a m ,otholllna

8 (7) Thallus orange )e1lo\\'. margtnallobule subcoO\e\ oriented in mJIJlIng manner, pon:' 10-14 : -IJln! (Fig 20)

Figure 20.-Thallu> of Ca/op/aca marina. Scale i; 1 mm.

. aloplaca marina

9 (4)

9 (4)

10 (9)

10 (9)

11 (9)

Spores one-celled ......................................................... : ............ . ...... .. .......... 11

Spores more than one-celled ............... '. ' .............................. . .............. . ................ 10

Growing on thallus qf Verrucariae; paraphyses gelatinizing, spores 10-15 x 3 .5-5JLm with 4 locules and distinct cen-tral septum (Fig. 21) ..................................... . ................................ . Stigmidium marinum

Figure 21.-Spore of Stigmidium marillulll. Scale is lO/Lm.

On barnacles or rock; thallus scant to evidently lacking; paraphyses persistent, spores 9-20 x 4-7 JLm with two locules (Fig. 22) ....................... . ........ . ..... . .... . .. .. . . .... . ... ... . .. .. .. ... . .. . . .. . Arthopyrenia halodytes

Figure 22.-Spore of Arthopyrellia halodytes. CD Scale is IOl'm.

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

Figure 23.-a. Thallus with pegs. b. Thallus with ridges. Scale is 1 mm.

a b

11 (9) Thallus smooth; pegs and ridges lacking . . . . ..... . . .. . . . .. .... ... .. .. .... .... . ....................... . ........ 12

12 (11) Thallus areolate (divided into fields by fissures , Fig. 24) . . . . . . . . . . . .. . ..................... .. ..... . .. . .......... 15

Figure 24.-Areolate thallus. Scale is lmm.

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

Figure 25.-a. Continuous thallus. b. Discontinuous thallus. Scale is lmm.

/ ("

/'

a

OC3 c:Jb

13 (12) Perithec ia elevated, often shiny, dOll1e -~haped to hellll\phencal or .~(llllelill1e, pOlntcd (Fig 2h) .. .... 14

Figure 26. - Thullu, \I ith cle, ah'd perot hec la .

Scale i\ I mOl .

13 (12) Peritheciaubll1erged to sltghtl) elevated (Fig. 27). ~pnrc., 8-11

14 (13)

Figure 27. - Thallu, ,,,th 'UbOlH~.d In slightl) ele'D ted perothc"u . "'Cille" I moo.

Spore· 6-11 X 3-5jtll1 ..

14(13) Spore 18-22xS-9jtll1 .... , ........

4-5 lilll. thallu .. thl~~, green to gro.:enl~h-hl<ld;

/

. . Vt.·rrucarw IIlL/C{)jll

.J )

--------------~/ ~--

~0rrll( ana micro pora

~errurllrl/l tflC/rnla

15 (12) Areole ltght bro~ n on dark. bro\\ n prothallu\ (Fig 2 ): penlhecl<l hm\\ n, pore., thl~ ~ \\ albl (II 5 15-2 . (27) x (4) 6-8( 14)jtll1 .. .. ..... .... ~ 'I;'rru({trlj/ III1UlIIl(re\Ct 1/1

a

Figure 28 .-a. \ertical section of perit heci um . Scale i. 0.1 mm. b. Thallus of l'errucaria Inlernigres­cens. Scale is 1 mm. b

15 (12) Prothallus not evident; thallu proll1lnentl) fissured. penthecla bro~ n or black .. spores thm-\\ ailed. 9-12 )0., 5-6jtll1 (Fig. 29) ..................................... .. .. . . . . . . . . . . ~ errll('aria cellThocarpa

a

Figure 29.-a . Vertical section of perithecium. Scale is 0.1 mm. b. Thallus of Verrucaria ceuthocarpa . Scale is 0.1 mm .

16 (11) Thallus areolate (thallus divided into fields by fissures , Fig. 30) .. ....... .. ...... ..... ..... .. ....... .. . .. .......... 17

Figure 30.-Areolate thallus. Scale is 1 mm .

16 (11) Thallus not areolate .............................................. ........... ........ .. . ....... ........ ... . 20

17 (16) Areoles bordered by raised juga (Fig. 31) .............. ... .................... .. ............................... 18

Figure 31.-Areoles bordered by juga. Scale is 0.1 mm.

17 (16) Areoles not bordered by juga (Fig. 32) ........................................................................ 19

Figure 32.-Areoles "ithout juga . Scale is 0.1 mm.

11

17, P nthecla u uall~ Immer<;ed.o_[} Ie border frequentl~ raled. spore 9-1:! X 5-6Jlm: juga rarel) limited to borders of areal • u uall) cattered \\ Ithm areole <Fig. 33): thallu more tran lucent \\et than dr) ................... ~errLlcaria degelii

a

Figure 33.-a. Vertical ,ection of penthecium. Scale is 0.1 mm. b. Thallus of lerrucar;a degelii. Scale is 1 mm.

b

18 (/7/ Penthecla elevated. often craterous. juga formmg a retIculate pattern (Fig. 34): thallus opaque dry. translucent wet

. . . .. . ................................... ... . VerrLlcaria amphibia

I-

a f i)!urc .H.-a. Vertical sectinn of perithecium. Scale is 1 mm. b. Thallus of ler­rllCana amphibia. (Speclmcn illu,trated i, thid.er and stored longer than that in J i)! 3'1b.1 '1rale i, I mm

1 t1 Juga ppean'lg a peg' prolrudmg fmm thallu, and al,o to some extent on pentheCla; thallus nmose-areolate, u ua h bl ..... kt ll-brN\n Llnu opaque (wet or df')): penthecla often large. 03-0.7 mm diameter, immersed or

I .. t'd rtl) I p<.:li I n ing dbrupti). ,omctimc, depressed on lOp (Fig. 35), spores 10-:!0 X 7-10/lm ..... VerruClIriC/ inC/LIra

ur < mm b

a

lion or p<rllh lum. 'lUI. I 11.1 It I I mm.

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

a

Figure 36.-a. Vertical section of perithe­cium. Scale is 0.1 mm . b. Thallus of Verruca­ria erichsenii. Scale is 1 mm.

W (16) Perithecia immersed , spores 9-12 X 5-6JLm; juga appearing as long sharp black ridges on borders of thallus, some­times extending across thallus , fissures occasionally develop along juga and simulate areolation; thallus usually green to cream colored (Fig. 37) .. . .... ......................................................... Verrucaria degelii

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

Scale is 1 mm .

10 (16) Perithecia raised and prominent (Fig. 38) ............................... ...... ........................ ......... 21

Figure 38.-Thallus with prominent raised perithecia.

III (20) Juga inclosing lenticular areas giving appearance of ripples on water, continuing on perithecium to form reticular pattern. Perithecium elevated, often craterous (Fig. 39); spores 9-20 X 4-7.5JLm ....................... Verrucaria amphibia

a

Figure 39.-a. Vertical section of perithecium. Scale is I mm. b. Thallus of Verrucaria amphibia. (Thal­us thinner than Fig. 34b. Cracks developed in storage.) Scale is I mm.

III (20) Juga scattered, not inclosing lenticular areas ................................................................... 22

22 (21) Juga appearing as pegs to usuall y short riCIges, usuall y straIght bur somerrme crescen snapeu, urnemrn::n:;o·a---------­

lescing with perithecium at base ; thallus usually dark o li ve green but lighter and with fewer juga in shade; peri-thecia hemispherical , often shiny (Fig . 40) ; spores 6- 1 I x 3-5J.tm .................................. Verruca ria dilmarsica

a

Figure 40.-a. Vertical section of perithecium . Scale is 0. 1 111111. b . T hallus of Verrucaria dilmarsica. Scale is I mm.

b

22 (21) Juga broader, irregular, and sometimes branched (Fig. 41) ........................................................ 23

Figure 41 .-Thallus \\ ith broad , sometimes branched juga. Sca le is I mm .

23 (22) Juga conspicuous, highly irregu lar, frequently branched. raised or immersed and appeari ng as con picuous black spots in wet thallus, often merged with peri thecia; wet thallus usually brown to amber and translucent (Fig . 42 );

perithecia up to 0.3 mm diameter, spores 8-9 x 4.5-7J.tm .......................................... Verrucaria erichsenii

a

Figure 42 .-a. Vert ica l section of perithecium . Scale is 0. 1 mill. b . Thallus of Verrucaria erichsellii. Sca le is 1 mill. b

23 (22) Juga prominent , black, and shiny, much thicker than in V erichsenii, frequentl y broadened into th ick irregular plates, espec iall y at thallus margin ; thallus usually grass g reen but darker in the sun and depigments rapidly in shade or in storage; perithecia typically hemispherical to globular with fl attened tops but may become quite irregular, angular, or dissected (Fig. 43); spores 8- 10 X 4-6/Lm ...................................... Verrucaria'striatula

a

Figure ·n.-a. Ver tical section of perit heciu m. Scale is 0. 1 mm. b. T ha llus of Verrucaria striatula. Scale is I mm.

ANNOTATED LIST OF SPECIES

Where distributions were derived from the literature only, author and date are given . All other specimens were examined by he author. Where such specimens were collected by others, the col­ector's name is given followed by the identification of the herbar­Ium retaining the specimen. The remaining distributions are from he author's collections.

Genus Arthopyrenia

Mrthopyrenia halodytes (Ny!.) Arn. Ber. Bayer. Bot. Ges . 1: 122. 1891. Verrucaria halodytes Ny!. Mem. Soc . Sci . Nat. Cherbourg 5:212 . 1857 .

DESCRIPTION : Thallus epilithic (siliceous rock), endolithic (calcareous rock and hells), yellowish if epilithic, greyish or

lackish-brown if endolithic. Perithecium almost wholly immersed n substrate or sessile with black hemispherical to broadly conical

'nvolucrellum , 0.15 to 0 .5 mm diameter, excipulum colorless to pale brown. Spores eight per ascus , ovoid, two-celled, one cell usu­ally broader than the other, 9-20 x 4-7 .5/Lm .

Swinscow (1965) considered this to be the only littoral species fthe genus and treated A. sublitoralis (Leight.) Arn. , A. Joveolata . L. Sm . and A. gyalectoidea Knowles as synonyms and reported range of 10-20 X 5-IO/Lm in spore size . Richard C . Harris (1975)

reats thi species as cO.,a Pyrenocollema. Arrhopyrenia halodytes is often associated with shells or

alcareous rocks but may also be found on siliceous rock .

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

NORTHEASTERN AMERICAN DISTRIBUTION: NEW ERSEY: Cape May Co. CONNECTICUT: New London Co.

I HODE ISLAND: Newport Co. MASSACHUSETTS: Barnstable

Co., Plymouth Co., Essex Co. MAINE: Cumberland Co., Sagada­hoc Co. , Hancock Co., Washington Co. NOVA SCOTIA: Yar­mouth Co., Digby Co. , Halifax Co. , Victoria Co., Cape Breton Co. NEWFOUNDLAND : West Coast Section, Northern Penin­sula 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, citrine in shade, orbicular, irregular, or subeffuse , small convex lobes contig­uous or. in center of thallus , minutely granular or tuberculate , whit­ish 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/Lm.

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

GENERAL DISTRIBUTION : Norway, Sweden , Poland (Nor­din 1972) , Finland (Ra'sanen 1927) , Germany (Erichsen 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.

Caloplaca microthallina (Wedd.) Zahlbr. Cat. Lich. Univ. 7:247. 1931. Lecanora microthallina Wedd. Mem. Soc. Sci. Nat. Cher­bourg 19:276. 1875.

DESCRIPTION: Thallus citrine, 1-3 mm diameter, comprised of scattered lobes on the periphery, 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/km.

Caloplaca microthallina 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 Verrucaria maura 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, Den­mark (Nordin 1972) .

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

Caloplaca scopularis (Nyl.) Lett. Hedwegia 52 :242 . 1912 . Lecanora scopularis 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, con­vex, 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/km, isthmus ca. 0.3 the length of the spore.

Caloplaca scopularis 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 dis­tinquished 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 (Rasiinen 1927), Germany (Erichsen 1957), Japan (Nylander 1890).

NORTHEASTERN AMERICAN DISTRIBUTION: MAINE: Cumberland Co., Hancock Co. NOVA SCOTIA: Halifax Co., Vic­toria Co. NEWFOUNDLAND: West Coast Section, Avalon Sec­tion.

Caloplaca verruculifera (Vain.) Zahlbr. Cat. Lich. Univ. 7:272.1931. Placodium verruculifera Vainio in Med. om Gron. 30:313.1905.

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

Specimens of this species. previously unreported from North America, were Identified for the author by 1. Poelt. It is easily con­fused 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 grantii Magn. Ann. Cryptog. Exot. 5(1):21. 1932.

DESCRIPTION: Thallus of irregular whitish-grey large gran­ules, KOH + yellow, medulla with small crystals. Apothecia con­cave 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/km .

Little of 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 material 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 Sec­tion.

Genus Lichina

Lichina confinis (0. Mull.) Ag. Sp. Algar. 1: 105. 1821. Lichen Confinis O. Mull. Icon. PI. Daniaw 5:5. 1872.

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

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

GENERAL DISTRIBUTION: England (Ferry and Sheard 1969) , Wales (Fletcher 1973b) , Finland (Ra'sanen 1927), Germany (Degelius 1939), Italy (latta 1909-1911), Norway (Havaas MSC), Sweden (Degalius MSC).

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

Genus Stigmidium

Stigmidium marinum (Deak.) Swins. Lichenologist 3:55. 1965. Sagedoa marina Deak. Ann. Mag. Nat. Hist. 1. 12:40, pI. 4, f. 13 . 1954.

o Parasymbl nt on lanoral nucana

lally V. mucn a and V. mIcro pora. I lung a I Ible thallu

own Penthc 18 usually resembling that of host II hen . I

ImmeniCd on V.mucn a and hcml!>phcncal on V ml ro pnm In\

lu rellum black with pigment extend 109 IOto h lin C Ipulunl

pores eight per a!>Cu • two-celled with ea h ell ften dl\ Id d b

pseudosepta. upper cell u ually lightly wider than th In" [ 10 15 3.S-S"m The penthccla of Sflgmld,um are C8"I) conru cd ",th that f the

ho t. flgmid,um. however. ha t" 0 elled pore of 10· 1 5 -

"i"nl SwinM:ow (I 965)rcponed pon; 10 I 4· f)"m . The com·

binatlon S,igmldium mannum "II made h) S" In ow when he

united Arthopyrrmu marilla (Ikak ) L Sm and A Irp,muCl ( yl.) Arn . either of Ihe'ie pe Ie appear In the onh Ameman

checklist (Hale and ulhcrson 1970) nor did SWin CO" (196 )

ThiS. therefore . repre Cn!

ne" record for onh Amenla.

(,E ERAL DISTRIBUTION German). Finland . Ireland , Jer

C}, England (SWinSCOW 1965).

ORTH EASTER 1ERICA . · DISIRlHl ' I'I<> ' ~I /\I.:

Cumberland Co. , Hancod, (\1 • 'EW JI~ RSI~Y (),e,ln ( 0 1: \\ ·

OUNDLAND \alon Sed Ion

Genu .. ~errucaria

The ,PCC'IC" of thl~ g\:nu lorlll,'m "\\ hleh nw) be l o nlinll llll o r

orl11 ,cattaed pall'he" 1l111",e arenl,ltc' or di\ IJ d lilt I dl rete, re· lk, or Illhe, The thalli \ ;lr) III ttlld,I1(' fro llJ 20 to son/{m . li nd

11.1) hl' all11o,t l"IlIllpktd) tr,ln'p,lrcnt to 1'11Ilrcl) oP,l4ul' In \) l11e

pc"e'. Ihl' tr .. n'p~lrerll) (II the thallu, l1la) be enhan d h \\ CI

mg. \\ hill' lIther, rel11dln oPd4U~ \\ hcn \ht Th~ th Iii Illa) be 111\1\\lh or rnughel1l'd 11) dark pl1int' or nJgl" rh~ nd)! S l li ed

uga (Sanll""l11 1(39) , \ar) In kngth, \\IJlh , and Ihl nc I he

al"t Ilngln ,lnd "ntllgcn) 01 thc Juga" nlll "n Wil hut II "PI .11'> hal thc) onglnatc ut \If Ju,t hel \\ thc un.lcc (II Ihc th,tllu In the

uthM\ c pencnl"C, thc) dll nllt ntcnJ tll thc uh trJIC c pt a t Ih

dge, llt thc a~lllc~ Often a JI\{lndllln 1\ l11adc 111 kl') bet" e n

lint unJ nJgc fllrrlh III thl' Juga, hut II " ,001111 n Ih .. t lh POint

~ mcrel) pcak~ l'n inulO. PlcUIIU' llr hldJ~n ndgc

The pcnthcl'la III all III th\: Iitt\lrJl Vc'mJcdnJC , \\lIh \\hl h Ih

uthM h \\ llrt..cd, ha\ c a dar" in\ luc~lIum hi h 01;) ' p~ d

I \ r

rrrurona ampl"bw R I 17

I, m /tl/no( I

merged, 0.1-0. 15 mm wide. involuerellum u~ually brown. eXLIpu lum hyaline to dark below. spores hyaline. ovoid H 10 >( 'i Sinn

VerrLlcaria celllhocarpa is easdy confuseu with Velrun/ntl cleRe

/ii when either i~ modified by environment or when Its '>lructun.: IS obscured by epiphytic algae The elefts of ~ (elhocarpa may

become darkened and, on cur,ory e:l.alllination. may resemhk the dark ridged border of arcoles of II t/egclll Careful eX<.!l1l1natlon 1\

needed to observe the absence of riuges Conversely SOlm:tlmes the ridge are scant on a thallus of V £lege/II anu It may by 1111\ta"en for that of II ceuthocarp£l.

Perhap~ the greatest obstacle to correet luentifleall<ln is ohslur­ing of surface detail by eplph)tlc algae POl11Ons of questlonahle thalli should be nooded with water to ,often anu expand the algae , If details are stdl obscured, the fIooueu portion Lan he slruhheu with a small water color brush True riuge, Me f~ml) re\l,tant til such gentle abraSion and \\ III pei"'lst untd the thallus becol11es too oftened by the vater to malfltaln Its Ifltegnt)

Accordlflg to Sante,son" "e) (1939), the pnme ulagnostiL featurc 111

II dege/ii vs. V. cl'lIIhoc(lfpa is the presence 01 nugcs nn V t!(,~"'" The author finds that t\\O other feature, are qUile usL'lul. The be t

of the two IS tendency for natness to conca, It)' betw cen nuges III an areole of V £lege/II compared with the cnm e\nc" of arellie anu branches thereof m II celllhocarpa, ~('/'nt('ana ct'/lIho((/rl'(/ IS trul) glabrous. FUl1her. from the limited matenal in the author\ three North Amencan collections il abo appear, that the 1m oluL n.:llurn (II II ceuthocarpa tends to be bro\\ n \\ herea .. that of V elcgc/1/ IS lh tinctly black

GE ERAL DISTRlB TIO ;-"ol1hern EUfll!X 'pnsbergen . Bear Island. Q\'aya Zeml)a. Ibena. Benng trails. Grecnl.md. Kerguelen, Antarctica (Lamb 1953), Washington L S A (collccleU by Fink and determined a ~hl1l lInll lIlalira h) lahlbruL"ner MSC) .

ORTHEASTERN AMERICA' DISTRIBCTIO '. "'E~W­

FOUNDLAND. East Coast Section

Verrucaria degelti R. Sant Ar". Bot 29A (10);31 1939.

DESCRlPTIO : Thallus rimo e. formmg lobes or dl\crete are­oles, amber, brown or black when dry. amber to brow n \\ hen wet. prominent black ridges bordenng areoles and often pentheLla. thin to

thick (100-320 nun). Perithecia unken to promlncntl) ral~ed. 0. 1-0.25 mm diameter, excipulum hyaline Spore, h) al inC. o\old 9-13 >

5-6/-tm. Though published in 1939. thiS species was kno\\ n onl) from Nor­

way until the author's collection, thus there are few specimens and little study of ecology upon which to ba e conclUSIOns about em Iron­mental innuences on variations. It IS rea onable to asume. however. that the nature of variations is consistent with that seen in other Verru­cariae. If so, color would be lighter m shade form, than un form and ridges would be fewer and thinner in shade forms than on sun forms. This kind of variation is probably the cause of confuSIOn of some specimens of V. dege/ii with V. cewilocarpa . The identity of a robust specimen of V. degelii is beautifully clear. However, the ridges may become scant and the specimen trongly resembles the morpho­type of V. cellthocarpa with darkened cleft~ between areoles. A sec­ond source of confusion is the epiphytic growth of algae on thalli of either V. degelii or II cewhocarpa. A growth of algae on V.dege/ii can obscure ridges and produce the glabrous appearance of V. ceUlilo­

carpa. Such a growth can also produce a shade condition for the

und.:rlYlng thi.llius and redul:l: the growth of nuges, On the other hanu. a growth of uar" a Igac on V ( ('lIIhlllorpCl, c~rc('}i.llly w hen dry, rna) ohscurc the color <lnu glahrous n<lturc of the Ihallus :.md gl\(; the illUSion 01 nugl:s on the cdge of arcnlc\. Onc (;,lnflOI over ernpha'l/l: the I III po rtalKl' 01 u.:te<.:lIng and rCrT1nvlTlg ,uch eplphyllc algil(;'

'r he thallu\ of V dl'Xc/1I v",nes greatl) In thJc~ncs!> and transpar­enl'\ depending on age or cxtent 01 uc\'elopmcnl. It I alm() t alwilYs i.lreol"te With only one ~ nowll eXlcptJOn fn thilt COl c, a rather large <lrca of month. lontllllHIU\ thillius was unounucd h) a rJlsed hlack ndge and iJ erad., al"o hordereu h) a hlal:k r.med ndge, c. tended IIlwaru t()\\arU thc l'enter At the eugcs nf thiS patch nlthallus Wa~ Illllre thilllu\ UI\ lued Into typical ul\uete arc(lle w IIh Imllar hla " horu.:r, ,\1llre comlllonl) a )lIung thallus" thm, ne<.lrly tr.ln'parcnt, \Illollth ,md hrllw n to t.m It" ,JreoliJtc and 1'; oilen dl\ IUed Into lobe .. h;. a dendntl<.: p.!llern 01 gnl(I\C\ In sUlh.J thallus, thc gfr)(l\C~ <.Ire usu.Jll) h)allne

I he IIntogcn;. "I these lichens h",s not hecn MudlCd ,lIld mu t he Inferred Inlln ohscn,ltlon llf wh.!1 could a ei.l II) be the re ult 01

cllndllillns III gnl\\th J III a 'Ing It .Jppcill'S. h we\er, thi.lt older thalli thlc,",en ,md de\ elop dl rete areole . u u.llI) \\ IIh blad ruel'; and fn.:quentl) \\ Ilh hI.! ~ lin.: Cllnnectmg them \\ IIh the pcmhelJurn 11K' e are.J' 01 hla~"n~ then eem to prolJlerate under orne Ondl­lI(1ns to produ e unci ·,ated pattern r raised Juga , mellme the hl.ILk .Jre.1 e tend hcnc.lth the pcnth L1um nd somellme Il doc 11"1.

The pc nthellllin I (llnunonl) IInme~'d, with or \\ Ithout ral~'d

nJge .1Tl)uOO the 0 II Ie At lime th rt' 1 a om e art'a r.lIS~d around

the, "lillie The iluthor ha 'en pmrmnently r.used cOO\ e pt:nthl: 13

nn the 'klr11e thallu <J un~t:n p..:nthet'IJ. The uegR.'C 01 pcnthe aal elc\allon arrc<lr, tll he d \ilndhle charatt:r of the pc Ie .

Spore an.: (1\ OIU hut wmeume polTlted \\ hen ~oung ,-\ clear pot commnnl <llthough not .11\13) appear. In the entcfofthc )ounger

['1'1'1:' S.lTltt: on ( IlI)Y) n.:pol1ed the r<lnge of pore lie for I t'ml

(lIna elt ~dll a I D-I J x 5,6jlrn. \\ hleh I coml tent \\ IIh that nh en eu h~ the author

GE;'\ER L DI TRIB TIO. ' .• OR\\ Y ( <lntc on IY3Y)

• 'ORTHEA TERI' J\~IERI -\ o. ,\lA

DI TRIBL TIO CHL' ETT

,\1 L E

E ex Co Dlgb) Co. Habra\ Co E\\'-

t Coast eetlon '-orthern Pemmula eLllon. Ea't Coast SClllon. \ <lInn tllHln

~(ITIi Car/(/ dlll1/aTlI(,(1 Ench., Lhriften t\Jtunl I. s Vcrclfl chIc \\ig-Hobteln 22.90.1937

DESCRIPTIO:\ Thallus entire. 011\ aleou . \1 ith p.:g. to shon ndge. thin (20-50jlml. opaque to tran,lucent \\hen df). more translucent \\hcn wet Pemheclu .. light I) dome-shaped to globose (usuall) heml'>pherieal). 0 1-0 25 mm diameter. usuall) .,hln:: exclpulum h)allne to dark belO\\ pore~ 0\ Old to reniform. color­le\s . 6-11 x 3.S-6/-tm

The thallus of Verrllcaria dilJllllrsica IS usual I) oll\·aceou,. but the darkness \ anes \\ nh light exposure. In bnght light the thallus IS dark oli ve and I f gro\~ n In the shade n IS a \ery light 0111 e green. It usually IS rather transparent to translucent. The transparenc) i~

increased by wetting upon examination. Shol1. most I} straight ridge~ are characteri~lic on the thallus . These become reduced in size and frequency in the shade modification. In this reduced condi­tion it is easily confused with the shade modificatIOn of V. erich-

·1 (,lIil and e~peCially with V .lIrl(1/li/a \.\ hose color 1\ al,o \lmli;1r Rldge\ often merge WIth the penthella ;11 the b;1\e

The pemhecla are convex to heml\phem:al and u\uJIl) \hln) Darkne\s of the lower exclpulumls a hlghl, vanable factor. Gener­all y ,ome carbo naceou\ granule\ tend to extend below the In\olu­cre llum a nd may con tInue \omewhat throughout the lower involucrellum (sec the !..ey) . However, the speCIes \hould not be characten/ed a~ haVIng an In volucrellum that I, dar!.. belo\.\, a \ le\~ adopted by Ench,en (1957) .

GENERAL DISTRIB TION : Germany (Erich,en 1957). or­way (Santeswn MSC) .

NORTHEASTERN AMERICAN or TRIBUTION : MAl E Cumberland Co ., Sagadahoc Co., Han cock Co. RHOD E ISLAND : Newport Co. MASSACHUSETTS : Pl ymouth Co., Es ex Co. NEW HAMPSHIRE : Rockingham Co. OVA SCO­TIA : Yarmouth Co., Digby Co., Shelburne Co., HalIfax Co., VIC­toria C . NEWFOUNDLAND : West Coast Section, Northern Peninsu la Section , East Coast Section. Not prev iously reported from North America .

Verrucaria erichsellii Zsch. in Erichs. Verh . Bot. Verelns Provo Brandenburg 70: 192. 1928.

DESCRIPTION : Thallus entire or areolate, blac!.. to blac!..ish­brown dry, amber and more translucent when wet , roughened by rows of pegs or ridges ; ridges often branched, ometlmes sub­

ergcd in thallus and vis ible only when wet; thallus 30-70Jlm hick . Perithecia elevated, co nical to hemispherical. spreadi ng III

irregular pattern at base, often with pegs or ridges as on thallus, . 1-0 .3 mm diameter, excipulum hyaline below. Spores OVO Id , col­riess , 8-9 x 4 .5-7Jlm. Vermca ria erichsellii demonstrate considerab le thallus vana­

ion . It is usually considered to be clearly nmose nr areolate. T,PI ­ca ll y this is so but it is not uncommon to find a thallus of V.

rici1s£'llii continuous in an e peciall) wet em ironment , w here the hallus becomes thic!.. and ge latinous. When dry the thallus IS usu ­lIy blac!..ish-brown to black. A grey morphotype \.\ Ith a texture

like that of graphite was collected in the Narragansett 8a} area. When moistened during examination the thallus incn:a~ed In trans­

arency, one of the be ·t tests of que tionable thalli. When wetted he) re\ cal a pattern of blac!.. mar!..ings ly plcal of the pattern of 'uges u~uall) seen abme the surface. The thallus of I' ertc/twII/I

t) picall)' ha short. often branched Irregular ndges or rows of oinb. frequentl) contluent \~ Ith or continUing up 0\ er the pemhe­

' ia . Ridges are longer than \\ ide and rarel) very hIgh In e\treme IrCUI1l~tances ridges become hIgher and ,harper forming cusp, Jnd

. rcte~ . In other e\trcmes the) Illa) be I 111111el cd \\ ithin the thallu, 10 be rC\ealed onl) b) \\ ettlllg

The mllst consIstent character l)f the pentheelutll appeaf', tll be he Im:gular 'preadlng base \\ hlch seem, mo ... t prolll)uneed \\ hen le\\ cd from abo \ e a \\ et thallu, l)r \\ hell the thallu, has been eeled frl)m the roc!.. anu placed 011 a ,llde alld illullllnated from ell)\\ The dlJll1etef', l)1 peritheciJ tend w lall bet\\ een 0 I ,lIld 0 ,

11111 l,d1ac!..e ( 1934) and E- nch en ( \95") bl)th II,t J 'iX)re rang' ,)j c-

12 :; 7/(111 \\ hlch agree, \\ Ith the Juthl)r .... l)b,en Jltl)n

GF FR-\L Dl TRIBL TIO, l, 'r llJl1\ en,'h en \9.--), Bnt-I,h Cl)lumbl,1 (Oh" l)J1 \ I ' l. S\\ edcn ,In{ , HI I l '" J\

tona Cn.. apt: Brt:tnn Co eetlon. nnhern Pemn,ulJ Se~tlon, \\ Ion e~lIlln

l errucarlll 1Il/('/'l/iI:It'\(l1/1 (0:)1 ) J·mh \ern B , \I:rclll Pm\ Brandenburg 70 IY3 IY24 I (fT/III/I/(I (ltllllobn/a \<lr 1111 " Rre1Cl'1l.1 ) I In Brenner. \kddel ,JIld So f • una I I I nn 13'125 188n

DESCRIPTIO brow n prothallu\ Pentheclu onm 11, u1n\ c • n ~ .:I mill dl rn ter. excipulum h)ullne nr Intt:rl11llt enti) Jar!..l'n d bclo\\ Sf}<

hyalIne . (hId walleu, tu,Iform. (II )1 15 21 (~71

( 14)Jlm .

The brown prothallu .... thallu" Jl1d pentheLla arc dl lIn 1I\ I: Th thallus I, thIn 0) compan\nn \\ Ith nthl'r ur 01.11 form I

s indlcJteu Jh,nc, pore \\ere found III th orth Amencan collection, a' 1.lrge a, repnrtcu h\ 1 n h n hut

the) were oloateu anu of at) plcal ,h.lpe \1 (1 t free n \\I:re tn

the range of 15 2.~ n·S,I111 Endl\l'n (1457Ii.,ted /111 both lIttoral anu n01l1Ittmal, therelnre. It 1I1a\ he J urn J lu

facultati\ e In It ... hahlh ThIS peLle \I ted t nh localIty III North An erIca anu thl I thc fmt rc on! tmm / \menca,

GE ERAL 01. TRIB TIO. C,l'rlll,lm (/ nch en 1<)5

ORTHE'\SHR, A\I[RIC\ DJ<';JRlBl TI) [\\ FOL DL '\0 -\\alnn cd Ion

I ('rmcaria //IOl/ret \\'ahlenh 111 \<.h <;uppl \Ieth L h 19 I (

rounded oni al con picuou penthecla. 6. rounded. not impre sed

penthe lat p .and7 pore "ltha"ldthof7-IOmic.(IengthIO-19 or at mo t 20 mic.)"·

Sante on ontinue b) de cnbing even denation ~ ith \\ hich he a ociate, name of varietie or \\ hat he con Iders to be sy no­

n1 m

.. I With" hlte or light bro\\ n clear prothallus .. V. -;,schackeana Ench~ .

, With green-gra) or greenish black thallus. ·var. jumosocine­rea Vain:

3 With connected or triflingl) fissured thallus . . V scorina Wedd:

4. Thallus urface with many strongly conspicuous pegs .. V aracrina Wg:

5. a With perithecia almo t entirely sunken into the thallus . 'V malmet Sen:

b \\,ith hemi~pherical to nearl) spherical perithecia. ·var.

I'romitlLlla Vain: 6. With clearl) impressed. frequently somev,hat irregular

penthecium top~ . V rrachinodes Norm: . V hdyrenii Erich,:. 'var aracrillotdes Vain:. 'f. el'Oluta Th . Fr.'

(?7 Spores" ith a width of II-IS mic .. V finn ma rkica Zsch: )."

The author has seen all of these form in North America except" V finnmarkica:' A common variation re ults from frequent washing '" ith water fr m rain or ",aves which causes the thallu color and texture to be~ome obcured by epithallic algae which produce a

smooth greenish urface.

GE:\ERAL DISTRIBUTION: ovaya Zemyla (Lynge 1928). Finland (Rasanen 1927). Germany (Erichsen 1957) , Spain to Fin­land (Degellus 1935). Italy (latta 1909-1911), India (Awasthi 1965). Japan ( ylander 1890), Greenland, Iceland, Bear Island, ·pitsbergen. Iberia, Bering Strait, Fuegia, Patagonia, Chile, Falkland Islands, and e\\ Zealand (Lamb 1953). Norway (Sante -son M C). ~eden (Santesson MSC), Wales (Fletcher 1973a) , England (Ferry and heard 1969). Brillsh Columbia (Brodo M Cl.

'ORTHE TER AMERICAN DISTRIBUTION: MAINE: (Plm ' A) t\:OVA COTIA Yarmouth Co .. Digby Co .. Shelburne Co .. Halifax Co . Vlctona Co , Cape Breton Co. NEW­f OL' Dl \. D \\e t Coast eCllon orthern Penin ula Section. I a t ('''J\t elltnn. Avalon Section NEW BRUNSWICK : dmpobell~\

I( rTl/(llna ml(m por<l } I Ann LI '\;at. Bot. IV. 3 175 I 55

DE:. RIP 110 I hallu u\u .. tll} bn)\\ n to amber. ,Ollletlllle, grc n or 'fey. moolh thm _0 )() I'm (l)ntlnuou~ M In patches but

Iran lu nt to qUI[~ Iran p.lrI:nt \\ hen

wet. Erichsen (1957) recognized a var. laerel'irens a characterized by a leek green thallu which is Ie s transparent. The author has

also ob erved thi in mate rial frolll the southern part of its range in orth America as well as a dark grey thallus in the same area. The

dark grey thallu has a color and texture resembling graphite used in

pencils. Upon wetting. the dark color disappears . The perithecia 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 (Erich­sen 1957) . Erichsen (1957) also recognized var. jriesiaca as having

a smaller perithecia of only 0.1-1.15 mm diameter. The author measured 230 perithecia (10 each from 23 specimens) and found a range of 0.08-0.28 mm except for a si ngle 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. Perithec ium 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 pores , but this is not

noticeably different from V dirmarsica and V srriarula. Spore sizes have been reported as 7-12 X 4-5/Lm (Lamb 1953),7- 11 X 5-7/Lm (Zschacke 1925). and 7- 11 X 4-5/Lm (Erichsen 1957) . Erichsen (1957) also recognized var. mLicosLila Sands. with spores 5-9 X 4-9/Lm . The author's spore measurements gi'/e a range of 6-11 X 3-

5/Lm.

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

NORTHEASTERN AMERICAN DISTRIBUTION : MAINE : Cumberland Co., Sagadahoc Co., Hancock Co. MASSACHU­SETTS: Essex Co. , Plymouth Co., Ba rnstable Co. RHODE ISLAND: Newport Co. CONNECTICUT: New London Co. NEW

JERSEY: Ocean Co., Cape May Co. NOVA SCOTIA: Colche ter Co., Cumberland Co., Yarmouth Co., Digby Co., Shelburne Co., Victoria Co. NEWFOUNDLAND: We t Coast Section, Northern Pen in ula Section, East Coast Section .

VerrLlcaria /Ilucosa Wahlenb . in Ach. Suppl. Mcth. Lich. 23. 1803 .

DESCRIPTIO : Thallus gra~, grecn to blacki,h-green, smooth and tough. conllnuou~, often ~Ith necral crack, in herbarium , peci­mens, 130- J 50 /Lm thick, usual I} opaquc (wet or dry); juga ab~ent; prothallus, IS pre,ent, "hlw,h. PenthccIa submergcd to ~J l ghtl)

raised. ,omellme,,, ith prominent o,tlOle'> surrounded by chilllney, c"clpulum clear bellm. Imolucrellulll 0 05-0 2 mm diameter.

pores 'Impie, ('olorie,,-, u\uall) o\Old, 8-11 )< 4-5 /Lm Thallu, of thl' ,pccle, I' ea,t1) rccognlLed In the field or In the

hcrbanum In old herbanum 'pellmen, It I~ bnm n Upon dry Ing It de\cll p' cralk, that are stnctl) necral anlf::tch not to be confu,ed \\ Ith the rimo\e are(llaliOn t) plcal of ~ //Iil/lrti \\ hltl,h prothallu, I' nften ,cen around a thallu\ on ,lllooth rock

Pcnthella are normally lIu\h to ,ltghtl) fJiwd or ,unkell When npe. the In\lllucrellulll tend\ III C\ Jglnate ,l1me\\ hJt pre,enllng a

The Ihallu of V mUlnlO 1 qUltc a n:: I ft n 'I'll" (l r ru 10 algac, u~h a Hilt!, "brallfla nd! J,Iu ,II /lillI/a II I 'I

.ommonl) found m" an 0\ r thcr rT\I and "h n:: Ih pcmhc Iii of Ih 0\ crgru" n mah.:nal proJ Ilhrou 'h Ih thallu f I III/ a ausang a pOI nllal ou c of Illi 11.11.: nil II lIun SJlOrc~ of V I//II( osa re not dl\tlO 11\ an I.:lIh r hape or 1I.c n

e on (I liN) n::ported th' lie of pon::, III be hi 'hi} \ dndblc 7 I

4 JAI1l

p) ulldlil an:: qUit ~lJl1lmon on I' 1111/(1110 Ih.11I1 I h' appear it ,Itt nnged .... lIh hnm II all I f,,~ur .... Idcl) ~al\ercu or III dll tef\ II

n,'t uncommon 10 fllld mOllnd madc lip of Jggrl.: ated p cllIdla

Gb 1:R L I>ISl RIBl' 110, S .... cd 'n, Of\\<I), 11Il1,lIlu, I c lind, r.lroc hl.lI1d~. (,crlllall). Irelan I, I mn c ( ante un 191',1). nglill1d (h'rf) .lI1d She.lrd 1%'11. \V.tle, (1·1 t~her 1<,/7l.l) (,rcell

~1ll1. Slh(n.l. hI gl.l. Au\...I.lI1d 1,1.lIld . (ampr. II '~land. e .... ~l·al,lI1d (L.II11r. 1'1511, Bnll h Clllul11hla (Bm<i(l \lS( )

. OR'I HI:AS'II:RN ,\~lI:RICA:,\ DIS', RlBl I J();\ \IASSA HLJSI:I IS' b,e\ (II .. 1'1)l11l1l1lh CII :-';I:\\' II,\\II'SHIRI Ild(lngham (II MAL'I: (1Il1lheriand (0. S.1g.ldah «(I. lidll Ild (0. I.lI1coln Co (i\lelTll ~ISCl. W.I hlllgtoll ('0 (1\,\ (OlIA ('nil he,ler ( 0, (ul11r.erland II, Yarmouth ( 0 , Dlgh) ". Shelhllrrll' ell ,Haltl.1 C",. VI IOrl.! ClI , C,IJX! Brdon ( 0

E,WHH'NI>I,\NI> • \1,tl1l'rn "l'nlll 111.1 SClII'lIl, \-.1 t '0.1 I Cl'tH"l, A\.rlon SCllIOIl

Ilm"I/It/lIi""o/clhnI-.1I1ilcdnd •. \I\",lugl.12 ) n.'i 19\\

DI:SCRII'IION lh.11I1I brl\ll III h. ll'llttnllou 10 ,.HlIl\ not relllah: I'l'II(Ill'lI.1 hJIt(l\\ ""lIll' til heflll ph n~.". I ~ 0'; Illlll mad. l' llpuhllll h).dllle hI darJ.. hdO\\ ~P(lrc h).llm • I "1"1

4'11ll 'hI' .!lllhni \ I-.Ilnl\ kdg III till'; pll'te~ I 110111 d tI) th IllIllJ

hIll III .1\ Jlhlbk herb.lnum IlCuml'n, loll~dcd no no\\ n onl n'l11 Illllg "'.111,1. • ' Y 1 h(, O1.II(,l1al ,cen n: ~Illhkd I Ill/( rOlI'''''1 I ,II' 1I1"dl',.lhk dl'g' 'C 1 he I h.lllu \\.1 h \\ nl h Jnd patdl\ fh 'nth~'u.l, 1<1, Ihl' Illl"t p.lIt. \\('fC h.III()\\ dnm , H dn (19 I .1\(' () I. (l III II I .. ' the dl,lllll'('f III pcnthcuJ and Iho c c n b\ ll' .llIth", 1111 111111111 th" r IIIg,' I h .111 III h.II11 (l'n \I llf thl ' .... lll" I' thl' 'I " "lie." t'd 1>\ H dn (19".1 I __ lim I hl .llIlhor \\ J unahle t,l I and pore 1>ut BOlCh (I h I 'p(lrll'd th,' ran '1'.1 Ih _" () 10,1111

ORIHI \'ill R (IR" SUIl,)1

~\IIRI \ 1\\

1/1//((/11 I /ll/lfl/fll \\.Ihl'nh m \ 11 uppl I th I I" _

I. fl

<, IIU \ nih nn

MSC), British Columbia (Ohlsson MSC) , west central Canada (Bird 1970), Cuba (Montagne 1838-42), Greenland (Lynge 1932) , Novaya Zemlya (Lynge 1932) , France (Harmand 1905) , Germany (Erichsen 1957) , Nepal (Awasthi 1965), England (Ferry and

Sheard 1969) .

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

Xanfhoria elegans (Link) Th. Fr. Lich . Arc!. 69 . 1860. Lichen £'/£'­

galls Link. Ann . Naturges. 1:37. 1791.

DESCRIPTION: Thallus foliose, red orange to citrine, rising off of substrate at least at tips, KOH + purple. forming rosettes about 2-5 cm diameter, lobes nodular-convex, including tIps . ApothecJa orange yellow to red orange, about 1-2 mm broad . Spore colorle~s.

polarilocular, 10-13 x 5-8/Am . Upon cursory IIlspection this species may appear to be cru~tose

but it does have a developed lower cortex and lifts off the substmte 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. pariefina it is devoid of soredia.

GENERAL DISTRIBUTION: Washington (Howard 1950). Alaska (Krog 1968), South Dakota (Wetmore 1967), New MeXICO (Rudolph 1953), Arizona (Nash MSC), Colorado (Wetmore MSC)' Wyoming (Shushan MSC), Montana (lmshaug MSC). Iowa (Imshaug MSC), Michigan (1m haug MSC). Minne ota (FlIlk MSC) . Wisconsin (Malachowski MSC) , New York (Brodo MSC). Mexico (Beaman MSC), Manitoba (GillIs MSC). Northwest Terri­tory (Prest 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 DISTRIBUTIO : MAl E : Sagadahoc Co., Hancock Co. NOVA SCOTIA : Digby Co .. Shelburne Co., Halifax Co., Victoria Co., Cape Breton Co. NEW­FOUNDLAND : West Coast Section, Northern Penin ula Section .

Xallthoria pariefilla (L.) Th . Fr. Lich . Arct. 67 . 1860. Lichen parifillusL. Sp. PI. 1143 . 1753 .

DESCRIPTION : Thallus foliose, orange to citrine, radiate, often rosettelike, KOH + purple , lobes flat, wrinkled, ca. 1 mm broad, usually forked at the tips, soredia absent. Apothecia numer­ous, ca. 5 mm broad , plane to concave with thin thalline margin disappearing with age. Spores eight per ascus, polarilocular, color­less, 10-13 X 6-9JLm .

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), Nov­aya Zemlya (Lynge 1928), Sweden (Degelius 1935) , Finland (Riisanen 1927), Italy (latta 1909-1911), India (Awasthi 1965), Cuba (Montagne 1838-42) , Tenerife (Imshaug MSC). England (Ferry and Sheard 1969). Wales (Fletcher 1937b) , Spain (w. & c. Culberson MSC).

NORTHEASTERN AMERICAN DISTRIBUTION ' MAINE: Cumberland Co., Sagadahoc Co., Hancock o. MASSACHU SETTS: (Fi~her MSC) . NEW YORK: (Brodo 1968) OVA SCO T1A : Halifax Co., Yarmouth Co., Dlghy Co., Shelhurne Co

SELECTED BIBLIOGRAPHY

AWASI HI . [) [) 19(,5 Cal,doguc 01 Ih< lichen, Irol11 India I'«plli. ['lIk, Ian . and Ceylon

N"," Hcu"'g,a Buhcllc Suppl 17.lnr

BI RD. [) 11I7f1 Ke), 10 Ihe lilhen, 01 "'cSI lcnlral Canau" The HernMIUI11 . Del'

Bioi Un" (algllr~ I11ll11cogr. p 1,2

BRODO.I M IY6K The lichen, III l.on g 1,lanu ~c" Yo" ,\ ,cgelallonal "nu 11""'lll

"nal)"s ~ T SllIle \lu, SCI Sen l:Iuli ~lfI. ,JfJ I'

DI GfLlCS. (J

1'1)5 Da' 01 an"che clemenl de, ,Irllulh·und laun Ikchlen lI11rll 'on SI..lIn·

ulnll"en ALia Ph\logeogr Suel 7 I .. II

I'I)'} Ole llechlcn 'on norrll "al1l1n L pp,.llI Ln" Ar"kr II 1·205

ERICHSEI' . C F F. 11I57 Flechlen fiorll ,on ~(lrd"e'lueUI'lhIJnd (jU\IO' FI,chcr Verlag.

Siuligan. p I -111

HRR) B \\ . anu J \\ SH~ARD

IQ69 ZnniJllnn of \Urrallllorallll .. hl!"\ It" hCHC' .HlIunJ the DeJlc: Penln ul.J

Pembrol..e,hlre . F,cld IUd ,.I 167

FLETCHER. A 19Da The ecolog) 01 manne IlillOral) lichen, on ome rod) shrrc, of

Angle,c) I.lchcn"I,,~.'1 5 ,6 --100 I'iDb The cC<llog) 01 manlime (,uprllllllor.l) lichen, on ,,)JTlC roc hore,

01 Angle,c). Llchenol<lg"15 40 1·-122

HALE . \1 E . Jr. and \\ L C LBERSO\ 1970. .. lounh chcl·I..I"1 01 Ihe lilhen ,)1 Ihe wnllnenl.1 Lnlled 51. Ie, and

Canada Br) ologl'l 73 -199-543

HARM AND. L. J 1905 Lichen, de France Erlnal Pan, .I 609

HARRIS. R 1'175 Ala onomlc re\ I"on 01 Ihe genu, Anhop) ranla \Ia"al L.I (A,co-

m)eele,) In :-"onh Amen«l \lich Siale n" Ea,1 Lan"ng. )01 p

HEDRICK . J 1933 C\\ genera and 'pccle, of lichen, Irom Ihe herbanum 01 Bruee Fin,

I ~1) cologla 25 ,0:1·316

Ho\\ARD. G E 1950 lichen, of Ihe 'laiC of Wa,hlngilln L nl\ \\a,h Pre" Scallie. 191 p

l"'lSHAUG. H A 1957 Alpine lichen, of "C'lern Unlled Siale, and adpc<nl Canada I The

macrollchen, B,) 0Iog"160 177 ·272

JATTA .A 1909-1911 Flora IIalica lC)plOgama Pa" III LIChen, Rocca S Cas-

ciano SJab,limenlO Tlpografico L Cappelli .160 p

KROG . H 1968 The macrolichen, of Ala,," 'or Polann,1 SI..r 1-1-1. 180 P

LAMB. I M 1953 AnlarCIlC P) renocarp lichen, DI>co\ er) Rcp . 25: 1-30 195~ Lichen, of Cape Brewn hland. ",0\ a SCalia Annu Rep 'all \lu,

Can 132 239-313

LYNGE. B 19::!.8 Lichen!'! from 0\ a) a Zeml) a (c\cI of AC£lnnpora and

LecU/wwl Rep SCI Re,ulls am E\ped ",0\ a) a Zeml) a 1921. -13_ 299

P 1932 . Lichen, from nonh ea" Greenland. II ~llcrollchen' S~r S\albard

hha\el. 81 1- 143 . 1937 Lichen, from \\e,1 Greenland. collecled chicfl) b) Th M

Fne, Medd . Griilnl 118(8).225 P

MAGNUSSO . A H. 1932. Lichen, from "e'lern Nonhamenca. malnl) Wa,hlnglon and

Ala,ka Ann Cr)plllg. E,ol. 5(1) 16-38.

1940 . Lichen, from cenlal A"a . Rep . SCI. E\ped r\onh-\\ . Pr", China 13.

168 p.

MONTAGNE. J. F. C. 1838-1842 . BOlanique-Planle, Cellulalric, . /1/ R. de la Sagra. H"lOire ph)­

sique. pollilque el nalUrelic de I'ile de Cuba 8: 105-237 .

NORDI . I. 1972. Coloplaco sect. Gasparrinia i Nordeuropa. Uppsala. p. 1- 184 .

NYLANDER. W 1888 . Lichenes Novae Zelandiae. Pari s. p. 1- 156 .

1890 . Lichenes Japoniae . Pari s. p. 1- 122 .

POELT. J. 1969 . Bestimmungssch lussel Europaischer Flechten. J. Cramer. Lehre . p.

1-259.

RASANE . V. 1927 . Uber Flechtenstandorte und Flechtenvegetation im westlichen Nord­

finland. Ann. Soc . ZooI. Bot. Fenn . Vanamo 7: 1-202.

RUDOLPH . E. D. 1953 . A cont ribution to the lichen flora of Arizona and New Mexico Ann .

Mo. Bot. Gard . 40:63-72 .

SANTESSON. R. 19?9 . Amphibious pyrenolichens I. Ark. Bot. 29A( 10) : 1-67 .

SERVIT. M . 1954 . C~skoslovenskl:: lisenJnlKY celed: Verrucariaceae . Nakladatelstv'-

Ceskoslovenske Akademie Ved . Prana .

SWINSCOW. T. D. V.

1965 . The marine species of Arthopyrellio in the British Isles. Lichenologist 3:55-64.

TUCKERMAN. E.

1882 . A sy nopsis of the North American lichens : Part I.. comp risi ng the Por­

lIIeliaeei, Ciadolliei, and Coellogolliei . S.E . Cassino. Boston. 261 p . WADE. A. E.

1965 . The ge nus Caloplam Th . Fr. in the British Isles. Lichenologist 3: 1-

28 .

WEBER . W A.

1962. Environmental modification and the taxonomy of the crustose

lichens. Sven . Bot. Tidskr. 56:293-333.

WETMORE . C . M .

1967. Lichens of the Black Hill s of South Dakota and Wyoming. Publ . Mus .

Mich . State Univ .. BioI. Ser. 3:209-464.

ZSCHACKE. H .

1925 . Die mitteleuropaischen Verrucariaceen . IV. Hedwigia 65 :46-64 .

1934. Epigloeaceae. Verrucariaceae und Dermatocarpaceae . Rabenhorst 's

kryptogamen -f1ora I , part 9,695 p.

Arthopyrel1ia ha/oc/w('s .....

Ca/op/aca lIlarina . . . . .. ..

Ca/op/aca I'erruclllijera .. .

Ca/oplaca microthallilla .. .

Caloplaca scopu/oris . .... .

Lecanoro gral1tii ... .

Lichina cOl1finis .... .

Stigmidiulll lIlarillU/1/ .

Verrllcaria amphibw .

Verrucano ceurhocorpa ..

Verruca ria degelii .. . ...

SYSTEMATIC INDEX

1.2.3 . 9. 15 8. 15 8. It)

... H. 15. It) H. It)

... 2.7.16 . 2.6 9. It)

12. 11 . 17 10. 17 . 18 . 19

.1 . 12 . 1.1 . 18

Vl.'rrucaria dUmanlca

Verrucana I.'nc/Jlel/II

Verrucano 11111.'1I1I1<r£' 1 ((' 11 1

VI.' r rll C(I r/{{ IIUlII ra

Verrucano IIl1crOl(lOI'({

Verrucano I/IUC().\{/

Verrllcana 11/ICIco/a

Verrucar/({ Ifr/(/Ilt/a

Xli/II/lima ("(mde/ar/{{

Xal/lhorw c/l.'gClIl\ .

XCllllhor/{{ (lctrll.'lino .

.. 2,3.14, IH. 20. 21 2, 1,13,14,18. 19.21

10, 17. 19 . 3.10.12,15.16.17,19. 20

3. 10. 17,20.21 . 2,4.10,17,20.21

10. 17,21 2,4,15.19.20,21

... 6,21

. .. 7.22 . ... 7, 22

ACKNOWLEDGMENTS

Preparation of the "Marine Flora and Fauna of the Northeastern United

States" is being coordinated by the following Board :

Coordinating Editor:

Editorial Advisers :

Melbourne R. Caniker, College of Marine Studies,

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

(Fonnerly with Gray Museum, Marine Biologi­

cal Laboratory, 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, Sharon, Mass. Ruth D. Thmer, Museum of Comparative Zoology,

Harvard University, Cambridge, Mass.

Roland L WIgle). Retired. atlonal Manne FI h eries ServIce. Northea~t FI~hcnc\ ('cntd, NOAA. Woods Hole, Ma~~

Robert T. Wllce. Department of Botany. nl\er sity of Massachuseth. Amherst . Mass

The Board e tablished the format for the "Manne Flora and Fauna oJ the Northeastern United States." inVItes systematists to collaborate \0 the preparation of manuals. reviews manu,cripts. and adVIses the Scit:ntlfic Editor of the National Marine Fisheries Service .

Thanks are expressed to Henry A . Imshaug for hIS a~sl'tance In research and advice in preparing the manuscript. Thanks an. al 0 expreS\cd to Melbourne R. Carriker for continued assistance \0 later research stagcs and manuscript preparation . Richard D. Wood was very helpful tn the field work in the Narragansett Bay area. Helen S. Taylor. Bnan M Taylor. and She ryl M. Taylor contributed both in the research for and preparallon of the mauscript. Adele Deni on and Ralph Gorton aIded tn the preparation of the

final manuscript.

COORDINATING EDITOR'S COMMENTS

Publication of the "Marine Flora and Fauna of the Northeastern United States" is most timely in view of the growing univeral emphasis on environ­mental work and the urgent need for more precise and complete identifica­tion of coastal organisms than has been available. It is mandatory, wherever possible, that organisms be identified accurately to species. Accurate scien­tific names unlock the great quantities of biological information stored in libraries, obviate duplication of research already done, and often make pos­sible prediction of anributes of organisms that have been inadequately stud­ied.

Ronald Taylor started his research on lichens in 1964 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 Editorial Board of the "Manne Flora and Fauna of the Northeastern United States ." Work on the "Marine Flora and Fauna of the Northeastern United States" by the Coordinating

Marine Flora and Fauna of the Northeastern United States :

COOK, DAVID G., and RALPH O.BRINKHURST. Annelida : Oligochaeta BORROR, ARTHUR C. Protozoa : Ciliophora . MOUL, EDWIN T. Higher Plants of the Marine Fringe. McCLOSKEY, LAWRENCE R. Pycnogonida . MANNING, RAYMOND B. Crustacea : Stomatopoda . WILLIAMS, AUSTON B. Crustacea : Decapoda. POLLOCK, LELAND W. Tardigrada .

LARSON , RONALD 1. Cnidaria : Scyphozoa.

Editor is supported by the College of Marine Studies , Univer~ity of Dela­

ware.

Manuals are available from the following: Superintendent of Documents, U.S. Government Pnnting Office, Washing­

ton, DC 20402, for a charge. User Service Branch, Library and Information Services, Division D822 ,

Washington Science Center, Building 4. Rockville, MD 20852. at no

charge as long as the supply lasts. ational Technical Information Services, U.S. Department of Commerce. 5285 Port Royal Road, Springfield, VA 22161, either as paper copy or

microfiche. for a charge.

Manuals are not copynghted. and so may be photocopied from the OAA Technical Report NMFS Circulars available in most major libraries.

The manuals so far published in the NOAA Technical Report NMFS CIr­cular series are lIsted below by author, title. circular number. and TIS

acce sion number.

Circular o. NTIS O.

374 COM 73 50670 378 COM 73 50888 384 COM 74 50019 386 COM 74 50014 387 COM 7450487 389 COM 74 51194 394 PB 257 987 397 PB 261839

CAVALIERE, A. R . Higher Fungi : Ascomycetes , Deuteromycetes, and BasIdiomycetes. COULL, BRUCE C. Copepoda : Harpacticoida.

398 399

PB 268 036 PB 268 714

CUTLER, EDWARD B. Sipuncula . PAWSON , DAVID L. Echinodermata : Holothuroidea . HO, JU-SHEY. Copepoda : Lernaeopodidae and Sphyriidae . HO, JU-SHEY. Copepoda : Cyclopoids Parasitic on Fishes . CRESSEY, ROGER F. Crustacea : Branchiura . BOVEE, EUGENE C ., and THOMAS K. SAWYER. Protozoa : Sarcodina : Amoebae. WATLING , LES. Crustacea : Cumacea . ZULLO, VICTOR A. Arthropoda: Cirripedia . CAIRNS, STEPHEN D. Cnidaria: Scleractinia . TODD, RUTH , and DORIS LOW. Protozoa : Sarcodina : Benthic Foraminifera . BUSH, LOUISE F. Turbellaria: Acoela and Nemertodermatida. TAYLOR, RONALD M. Lichens (Ascomycetes) of the Intertidal Region .

403 PB 273 062 405 PB 274 999 406 PB 280040 409 PB 281969 413 PB 222 923 419 PB 285 538 4:23 PB 296460 425 PB 297 676 438 PB 124520 439 PB 225 053 440 PB 219 387