The Lichens of Fakahatchee Strand Preserve State Park, Florida : proceedings from the 18th Tuckerman Workshop

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The Lichens of Fakahatchee Strand Preserve State Park, Florida : proceedings from the 18th Tuckerman Workshop
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Lucking, Robert
Seavey, Frederick
Common, Ralph S.
Beeching, Sean Q.
Breuss, Othmar
Buck, William R.
Crane, Lee
Hodges, Malcolm
Hodkinson, Brendan P.
Lay, Elisabeth
Lendemer, James C.
McMullin, Troy
Mercado-Diaz, Joel A.
Nelsen, Matthew P.
Plata, Emily Rivas
Safranek, William
Sanders, William B.
Schaefer, Harold P. Jr.
Seavey, Jean
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Florida Museum of Natural History
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THE LICHENS OF FAKAHATCHEE STRAND PRESERVE STATE PARK,
FLORIDA: PROCEEDINGS FROM THE 18t TUCKERMAN WORKSHOP


Robert Lucking,'20, Frederick Seavey2, Ralph S. Common3, Sean Q. Beeching4,
Othmar Breuss5, William R. Buck6, Lee Crane7, Malcolm Hodges8, Brendan
P. Hodkinson9, Elisabeth Lay10, James C. Lendemer", R. Troy McMullin'2,
Joel A. Mercado-Diaz13, Matthew P. Nelsen1,14, Eimy Rivas Plata' 15, William
Safranek16, William B. Sanders17, Harold P. Schaefer Jr."8, and Jean Seavey19


ABSTRACT

Fakahatchee Strand Preserve State Park is located in Collier County at the extreme southwestern corner
of Florida, close to Everglades National Park and Big Cypress National Preserve. The 18tL Tuckerman
Workshop, an annual gathering of professional and amateur lichenologists and mycologists from the
United States and Canada, this time with additional participants from Puerto Rico, Peru, and Austria, was
held at this locality from March 1-7, 2009. Lichens were collected over a five day span from four sites
within the Preserve. Together with previously made collections, the survey produced a total of 432 taxa, 18
of which are new to science and 89 are additions to the North America checklist, six of which are also new
to the New World. The new species are: C. ,!, -' ,Jii.r floridana Hodges & I .n.1 i / G . ', ,. '1*., i
Common & Lacking, C).', -,'1, -,' miniaia Vain. ex Lucking, Di)., *.,i microsporum M. Ciceres &
Lacking (formally described in a separate paper), FIissurina (v. -.-',:.,1, ,i r Common & Lucking, Fissurina
analphabetica Common & Lacking, Fi'ssurina confusa Common & Licking, Fissurina inspersa Common
& Lacking, Fissurina pseudostromatica Lacking & Rivas Plata, tissurina subcomparimuralis Common
& Lacking (formally described in a separate paper), Fissurina fuckermaniana Common & Lacking,
Fi'ssurina varieseptata Common & Lucking, Graphis .1q, , ?,/', 4.,,,, Common & Lacking, Ifl,. !., i.r
., /,, ,.a , Common & Licking (formally described in a separate paper), Heiomasia seaveyorum M.
P. Nelsen & Lacking (formally described in a separate paper), Ph'i.. ;, .r !,i.; delicatula Common &
Licking, lapellaria floridensis Common & Lacking, and A ',IA //,./ i rgramulosa Lacking & Rivas Plata.
Further, the following three new combinations are proposed: Chapsa platycarpoides (Tuck.) Breuss &

'Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605 2496 USA; 'Dan Beard F 1. I I.: -.. .verglades National
Park, Homestead, FL 33030 USA < natureguides@mindsping.com>; 3534 Fenton St., Lansing MI 48910 USA < common@msu.edu>; 4301 Connecticut
Ave., Atlanta, GA 30307 USA ; 'Naturhistorisches Museum Wien, Botanische Abteilung, Burgring 7, 1010 Wien, Austria bg9.at>; 'Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458 USA ; Illinois Natural History Museum,
Natural Resources Bldg., 607 E. Peabody Dr., Champaign, IL 61820 USA < leecrane@inhs.uiuc.edu>; 'The Nature Conservancy in Georgia, 1330 West
Peachtree Street, Suite 410, Atlanta, GA 30309 USA < mhodges@tnc.org>; 9Department of Biology, Box 90338, Duke University, Durham, NC 27708, USA
< brendan.hodkinson@duke.edu>; 10239 Marlborough St., Boston, MA 02116, USA < elisabethlay@comcast.net>; "Institute of Systematic Botany, The New
York Botanical Garden, Bronx, NY 10458, USA ; "Biodiversity Institute of Ontario Herbarium, Department of Integrative Biology,
University of Guelph, Guelph, ON N1G 2W1, Canada ; "International Institute of Tropical Forestry (USFS), Jardin Botanico Sur,
1201 Calle Ceiba, San Juan, PR 00926, Puerto Rico ; "Committee on Evolutionary Biology, University of Chicago, 1025 E. 57h
Street, Chicago, IL 60637 USA ; "'Department of Biological Sciences, University of Illinois-Chicago, 845 West Taylor Street (MC
066), Chicago IL 60607 USA < erivasplata@ fieldnuseum.org>; 16Department of Molecular Biology and Microbiology, College of Medicine, University of
Central Florida, Orlando, FL 32816, USA ; "Department of Biological Sciences, Florida Gulf Coast Universit.y Ft. Myers, FL 33965,
USA; '25 AMedway Street, Dorchester, MA 02124, USA ; "Dan Beard Research Building, Everglades National
Park, Homestead, FL33030, USA j Seavey@nps.gov>; 0Corresponding author <, i.. i :~ i .....a .. >; authors after Licking, Seavey, and Common,
are in alphabetical order.

Clicking, R., Seavey, F., R. S. Common, S. Q. Beeching, O. Breuss, W. R. Buck, L. Crane, M. Hodges, B. P. Hodkinson, E. Lay, J. C. Lendemer, R. T. McMullin,
J. A. Mercado-Diaz, M. P. Nelsen, E. Rivas Plata, W. Safranek, W. B. Sanders, H. P. Schaefer Jr. & J. Seavey 2011. The lichens of Fakahatchee Strand
Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


I iil int Fissurina intercludens (Nyl.) Lucking & Rivas Plata, and Fissurina mexicana (Zahlbr.) Licking
& Rivas Plata. Six -.,1-e-. are for the first time reported for the New World: i; l.: ,:'.t interveniens Nyl.,
I! ,i'i,;ij. .i ,/i i~uL Nyl., C il, I/ *. r ii' (Hale) Mangold, Fissurina crassilabra Mont. & Bosch,
Stirtonia dubia A. L. Sm., and Stirtonia macrocarpa Makhija & Patw. Further 83 species are additions to
the North American lichen checklist: 1i Ji n. i ,. & endachroa (Malme) Marbach, Anisomeridium subnexum
(Nyl.) R. C. Harris, !.: ,l '.i ,uin ,,ll, (Fee) Nyl., I-:., .. 1,i i/n cinerascens Vain., I !,,;/i. li,,t,
geminiparum (Malme) R. 'S.,i i- i ,. Iii /, ', ../m li /!, P,,.LOcl I1, 'A.h. /imuJ Ii/, l .M ll.
Arg., Byssoloma chlorinum (Vain.) Zahlbr., C: ,!..,J ., editae V6zda ex Chaves & I i,-1 i,,. CI I,D .,'i.;
lecanorella (Nyl.) Kalb & V6zda, Calopadia .i ',,.ii/'.i (Nyl.) V6zda, C.'1 .,.nhli subcoerulescens
(Zahlbr.) V6zda, (C '. 'i." chionostoma (Nyl.) Rivas Plata & Ml.-r-,-cl,1 Chapsa platycarpoides (Tuck.)
Breuss & I iJ"l Itn C i ,'inmi congense C. W. Dodge, Coenogonium geralense (P. Henn) I i*kl int
C,',.-ii~'',.!i!te luteocitrinum Rivas Plata, Licking & Umafa., Coenogonium subdentatum (V6zda &
G. Thor) Rivas Plata, I i, 1 Umana & Chaves., Coenogonium '-.;j;ilVi .,iVw (V6zda & Farkas)
Li,-l n,. Aptroot & i;". Cn I ;i ,",::. Iwilmsiana (Mill. Arg.) Grube, Cryptolechia nana (Tuck.) D.
Hawksw. & Dibben, dCj 'i., 'i, ii i'.. i ,i il Arg.) R. Sant., Cir i~"' i. i, punctosorediata Sparrius.,
Dictyonema pi' /I ',.. "u'" i I ll Arg.) Zahlbr., Dictyonema sericeum f.- i! i/7i ,,','i::': Parm., Echinoplaca
leucotrichoides (Mull. Arg.) R. Sant., F.l. ,ic I.e leucocheila (Tuck.) Ii6l iu e Serus. & Kalb, Fissurina
11: ,., i1:,i, (Nyl.) 2 1.,'1 e Fissurina egena (Nyl.) Nyl., Fissurina mexicana (Zahlbr.) Licking & Rivas Plata,
Fissurina radiata Mont., Fissurina /i,. I.I,. !/ /a (Nyl.) Staiger, CA, P/'l-. if., I fi-f'c (Mull. Arg.) I ikl ir
Graphics i, ,:t. ;. Lacking & Umafia., Graphis assimilis Nyl., Graphis caesiocarpa Redinger, Graphis
caribica I itJ in, G a i/, .*if!. Zenker., CG oJl cupei Vain. ex I iil in_ Graphis ,i '-.. i, r' Nyl.,
Graphis handelii Zahlbr., Graphis /. ':u,,., Kremp., Graphis oshioi M. Nakan., Graphis pseudocinerea
I ikJ iw_ Graphis sauroidea Leight., GiC,,i' .':/i-,.Uii M. Caceres & I ikJ inw' Graphis subflexibilis
Lacking & Chaves., Graphis \ ,,ii,.' i' ':., Mull. Arg., Gyalectidium ,!i' ... Herrera-C.-in'' p,--. & I ;i-l ni.
Licking & G. Thor, H.. i p,,. ihil, echinatum Aptroot, LOcking & Will-Wolf, Lecanora achroa Nyl.,
Lecanora elapheia Stizenb., Leucodecton i :..p:'n. ii::.: (Nyl.) A. Frisch., Malmidea j, .. !, if ',iiN
Arg.) Kalb & Lik1 inu. Mahnidea gyalectoides (Vain.) Kalb & I ii i. MIalmidea leptoloma (Mill.
Arg.) Kalb & IT I-1 ;,in. Malmideapiperis (Spreng.) Kalb, Rivas Plata & Lumbsch, Malmidea rh..;d,-pi-:
(Tuck.) Kalb, Rivas Plata & Lumbsch, Mahnideao i. ., i. -. Kalb, Malmidea vinosa (T--lJ,..) Kalb, Rivas
Plata & Lumbsch, Myriotrenma ,i, 4 '; //" ,,h (Nyl.) Hale, 0O !i/'/. ,"'. auberianoides (Nyl.) Mill. Arg.,
Ocellularia obturascens (Nyl.) Hale, Pertusaria iH ,,:i/, ; i, i T., I,i Dibben, ,i., '..! / / i,' flavescens Dal
Forno & Eliasaro., Ph!.. ':'..;i' i:. i ,,,.! .i. ., (Fee) Mull. Arg., FTi .. 1'i. ,i5,. leiogrammodes (Kremp.)
M ull.Arg., 1 'li h... .'i .. ~ /n. i.., 'i (Kremp.) I idl I I J .. ,u ,i,'.--. nylanderi (Vain.) Zahlbr.,'/ ,i, ..-**. Ii/i,'
scalpturata(Ach.) 2i.-D, Phaeographis schizolonma(Mill.Arg.) Mll. Arg., P/i //.1 ,/'., i., ilacerataTimdal,
Pseudopyrenula i ,.-- .,.', ,M / Mill. Arg., Pseudopyrenula subnudata Mill. Arg., P-".i .. ,'i, 'u costaricensis
Henssen, Pyrenula brunnea Fee, Pyrenula sexlocularis (Nyl.) Mull. Arg., Y ';l, ';-,,,miiu marginatum
Licking & Lumbsch, Strigula orbicularis Fr., ./.' i :l schizospora R. Sant., Ti.' 1luH i .i ~li' 1II..-, I '!ii
L I-l i,-, /. ,. ii, /1 i 1amalmei R. Sant., and Thelotremap. in. b.''i:'i iif Nyl. The high number of species found
within a relatively small area, which corresponds to almost 10% of all lichens currently known in North
America, i. p i I iSi, i-, perspective by comparing it with other protected areas in the United States. It is explained
by the high carrying capacity of (sub-)tropical vegetation for epiphytic and particularly crustose lichens.
Keytables and image plates are presented to facilitate the identification of '.-e,;- -. in larger crustose genera.

Key Words: Ascomycota; lichens; new species; biodiversity; biogeography; Florida






LUCKING ET AL.: Lichens of Fakahatchee t.i i Preserve State Park


TABLE OF CONTENTS

In tro d u c tio n .......................................................................................................... 12 9
M materials and M ethods................................................................ ............. 132
R results and D discussion ................................................ ............................ 137
New or Otherwise Interesting Species............................................ .............138
C i.1, j, iJ;,floridana Hodges & LuJ,_.iil new species................................ 138
Calopadia .ii.I..-:,/ Common & Licking, new species ...........................139
Chapsa platycatpoides (Tuck.) Breuss & Lucking, new combination.......... 140
Cryptothecia miniata Vain. ex Lucking, new species................................ 140
Fissurina ,.i_:-.. -it, L Common & Lu.l.:_. new *T{*. '-;-.. ....... .............. 141
Fissurina analphabetica Common & Lucking, new species........................ 142
Fissurina confusa Common & Lucking, new species .................................. 142
Fissurina i ,/,. i ., Common & Licking, new ..f ,-i- ............ ................. 143
Fissurina mexicana (Zahlbr.) L ui,_l.1i & Rivas Plata, new combination
(with remarks on Fissurina intercludens (Nyl.) Licking & Rivas
Plata, new com bination)....................................................... .............. 144
Fissurina pseudostromatica Lucking & Rivas Plata, new species ............... 145
Fissurina luckermaniana Common & LuJl.Ai~. new species...................... 145
Fissurina varieseptata Common & Lucking, new species......................... 146
Graphics .. i ,. i.. i .- i ..i Common & L l. iii new species ......................... 147
/',. i ,,,',iii delicatula Common & Licking, new species......................... 148
Tapellaria floridensis Common & Lucking, new species........................... 149
7.'T. I/. / i., granulosa Lucking & Rivas Plata, new species........................ 149
Annotated Checklist of Lichen Species from Fakahatchee Strand
Preserve State Park ................. .. .......... ........ .......... ................ .......... 150
Acknowledgements .................. ................ .............. .... ............... 178
Literature Cited .................................. ..................... ... 178

INTRODUCTION
tropics (Aptroot 1997; Aptroot & ; .-i, 1997;
Lichens have long been believed to be most Sipman & Aptroot 2001; Feuerer & Hawksworth
abundant and speciose in cool-temperate areas, a ''*1-7; Lucking et al. ''^'). Lacking et al. '-')
notion supported by a recently published survey estimated that about half of the predicted global
from Alaska and its global comparison with other 26,000-28,000 lichen species occur in the tropics,
published inventories (Spribille et al. 2010). even considering that the tropics cover less than
However, these figures are misleading, as they 25% of the Earth's land surface. Our ongoing
compare areas of very different size (of more surveys in the Neotropics, particularly in Mexico
than two orders of magnitude) and also include (Los Tuxtlas Biosphere Reserve), Costa Rica
heterogeneous accounts of lichenicolous fungi. At (Las Cruces and La Selva Biological Stations,
smaller scales, e.g. looking at uniform area sizes Corcovado National P.-1 i. Venezuela (Henry
of 100 km 10 km 1 km2, or even one hectare, Pittier National Park), and Peru (Los Amigos
lichen species richness clearly increases towards Biological Station) I i numbers of 500-600 or
lower latitudes, like that of many other organisms, more species per km2 of rain forest vegetation, about
a fact realized only recently. This is particularly half of them foliicolous (Lucking 1999; Herrera-
true for crustose epiphytic microlichens which, for Campos et al. 2004; I iiJ i;tn unpubl. data). In
many groups, have their highest diversity in the quantitative studies, inventories revealed totals of






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


approximately 150 corticolous lichen species on 50
trees in the forest understory (Caceres et al. 2008),
as many as 250 corticolous .f-pri-;i, on less than ten
trees in the forest canopy (Komposch & Hafellner
1999, 2000), as many as 173 corticolous species on
a single tree from top to bottom (Aptroot 1997) and
as many as 300 foliicolous species on 100 trees and
shrubs (Lul.ilkg 1999); as many as 49 foliicolous
species were found on a single leaf (Lucking &
Matzer 2001).
Extratropical sites with similarly high or
higher species numbers (excluding lichenicolous
fungi) lr,-i in Spribille et al. (2010) are either
very large in size, such as Cevennes National
Park (France) with 973 lichen species (2297 km2),
the Bavarian Forest (Germany) with 841 species
(over 9000 km2), Pechoro-Ilychiskiy Nature
Reserve (Russia) with 790 species (7213 ].ill),
the Gurktaler Alps (Austria) with 774 species
(2950 ].ill), the d'Aosta Valley (Italy) ..iill 631
species (3262 km2), and Isle Royale National
Park (U.S.A.) with 596 p.....i-.-.. (2314 km2), or
include a heterogeneous and diverse geography,
such as Tatra National Park (Poland) with 864
species (211 km2), Berchtesgaden National Park
(Germany) with 813 species (210 km2), Klondike
Gold Rush National History Park (U.S.A.) with
668 species (53 km2), and Vega Island (! i .. .')
with 649 species (163 km2). None of these areas
maintain such large numbers at the small scale, but
rather they are the result of combining different
vegetation and substrate types with substantial beta
diversity. For example, the figure of 766 species
for the Klondike Gold Rush National History Park
(Spribille et al. 2010) is composed of 668 lichen
.-.....<-.. and 98 lichenicolous fungi, which were
sampled from two different units within the park.
The larger unit, Chilkoot Trail, harbored 510 lichen
species within an area of 35 km2. Considering that
these include lichens from all substrata (rock, soil,
bark, bryophytes), that number is substantially
lower than 500-600 *p ...i--. of lichens that can be
found solely epiphytic within 1 km2 of tropical rain
forest.
The high -.pfi-; richness in the tropics is
attributed to several different factors (Shmida &


Wilson 1985; Stocker et al. 1985; Linsenmair 1990),
including the absence of frost and the generally near-
optimal conditions for photosynthesis, the strategy
of 'escaping' competition by being rare and having
effective reproductive mechanisms, a tendency for
tropical communities to have a high level of entropy
or 'disorder' by dispersion of populations, high
dynamics with frequent intermediate disturbances
(Connell 1978; Huston 1979; Schupp 1992; Burkey
1994; Hubbell et al. 1999) which, along with
biological feed-back mechnisms (Janzen 1970;
Becker et al. 1985; Pirozynski & Hawksworth
1988; Armstrong 1989; Waterman & McKey 1989;
Wills et al. 1997) prevent competitive species from
becoming dominant and instead create a mosaic
of successional stages within communities, and
the notion that diversity begets diversity in an
'autocatalytic' fashion. Some of these hypotheses
have also been tested in tropical lichens (Lucking
& Bernecker-LUcking 21,,, "'1', LuI.Mll 2008).
The continental United States covers an
area ranging from the Arctic in the north to the
subtropical deserts and subtropical coastal swamp
forests in the south. Accordingly, the number of
lichen species estimated to occur within this area
is large, possibly close to 5,000 -. ...- --. Most areas
and ecosystems are comparatively well-studied
with regard to their lichen biota (Harris 1990, 1995;
Bennett & Wetmore 1999, 2005a, b; Brodo et al.
2001; Bennett 2006; Thomson 2003; Esslinger
2010), especially the Southwest and specifically the
Greater Sonoran Desert Region, with about 1,800
species included in a recent monographic treatment
(Nash et al. 2002, 2004, 2007). Other lichen-rich
regions that have been partially inventoried are
the Ozarks and part of the Appalachians in eastern
North America (e.g. Hale 1957; Dey 1978; Peck et
al. 2004; Harris & Ladd 2007, 2008; Knudsen et
al. 2007; Amtoft et al. 2008; Harris 2009; Harris &
Lendemer 2009; Lendemer 2 ,ic'. b, c).
The chiefly wet, (sub-)tropical parts of North
America are mostly restricted to the peninsula
of Florida extending westward along the coast
through Alabama, 1i;-- *--i 'i. Louisiana, and
Texas. But some usually tropical elements that
can tolerate more temperate conditions, including






LUCKING ET AL.: Lichens of Fakahatchee Strand Preserve State Park


some foliicolous lichens, also extend northward
into coastal Georgia and the Carolinas and in some
cases the Delmarva Peninsula and southern New
Jersey (Licking et al. 2007; Lendemer & Knapp
2007; Lendemer & Yahr 2004). Recently disjunct
populations of typically tropical taxa have also
been uncovered at middle and low elelvations of
the southern Appalachian Mountains (Lendemer
2009; Lendemer & Tripp 2008, and unpubl. data).
Many treatments on tropical lichens are
available for these areas (Moore 1966, 1968;
Skorepa 1968; Reese & Tucker 1970; Serusiaux
1979; Tucker 1979, 1981; Tucker & Harris 1980;
Smith 1986; Harris 1987, 1988, 1990, 1995;
Thor 1988; Harris & Wheeler, 1988; Griffin et al.
1995; Lumbsch et al. 1995; Ekman 1996; Staiger
& Kalb 1999; Marbach 2000; Buck & Serusiaux
2000; Grube 2001), including the classic works
by Calkins (1885, 1886, 1889; Eckfeldt & Calkins
1887a, b), Maller Argoviensis (1895), Rolfs
(1901), Merrill (1913), Bouly de Lesdain (1933),
and Herre (1942). Despite the large amount of
lichen diversity documented in these works,


which span over a century, previously unknown
and unreported taxa continue to be found in the
region at a remarkable rate (Lucking & Caceres
2004; Safranek & Lacking 2005; Perlmutter 2006;
DeBolt et al. 2007; Lacking et al. 2007; Lendemer
& Knudsen 2008; Lendemer & Lumbsch 2008;
Grube & Lendemer 2009; Lendemer et al. 2009a-
d; Seavey 2009; Seavey & Seavey 2009; Tucker
2010; Nelsen et al. 2010; Tripp et al. 2010).
These papers document the extraordinary
diversity of the region's lichens, especially of
crustose epiphytic microlichens, suggesting that
many more species await discovery and could
provide substantial additions to the North American
Lichen Checklist (Esslinger 2010). With this in
mind, the 2009 Tuckerman workshop (Fig. 1), as
part of the annual lichen workshops organized by
the Eastern Lichen Network (http://www.nybg.
org/bsci/lichens/eln) was held at Fakahatchee
Strand Preserve State Park, which together with
Everglades National Park, Big Cypress National
Preserve and Collier-Seminole State Park, form
a large protected area covering the southern and


Figure 1. Group photograph of the 2009 Tuckerman workshop. Standing from left to right: Lee Crane, Sean Beeching,
Frederick Seavey, Malcolm Hodges, Richard Fagan, William Buck, Elisabeth Lay, Mike Owen, Harold Schaefer,
Matthew Nelsen, William Sanders, Troy McMullin, and Brendan Hodkinson. Kneeling from left to right: Jean Seavey,
Bob Nesmith, Alicia Campanella, James Lendemer, William Safranek, Ottmar Breuss, Joel Mercado-Diaz, Eimy
Rivas Plata, Natasha Licking Rivas Plata, and Robert Licking. Color version available online at Ihp \\nu\ Ilmnh
ufl.edu/bulletin/vol49no4/figuresl-2.pdf.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


southwestern part of Florida peninsula.
The workshop was organized as part of
an ongoing NSF workshop project towards
an inventory of neotropical crustose epiphytic
microlichens (Lucking et al. 2009; http://
www.nsf.gov/awardsearch/showAward.
do?AwardNumber=0715660) and served several
objectives: extending the area covered by the
annual Tuckerman workshops to southern Florida
and thus the most tropical part of the continental
United States; document in more detail the tropical
element of the lichen biota of the continental United
States; train participating colleagues and students
in tropical lichen taxonomy, with emphasis on
crustose epiphytic microlichens; and produce a
guide to crustose epiphytic microlichens found
in the area to be used by the park personnel and
ecotourists.


MATERIALS AND METHODS

STUDY AREA
Fakahatchee Strand P i.... -. State Park,
a narrow strip of land oriented north to south, is
located in Collier County, Florida, and is about
3-5 miles (4.5-8 km) wide and 20 miles (36 km)
long (Fig. 2). Hydrologically, it is a _..,1_,.i- and
narrow drainage basin channeling the waters from
low-lying terrains to the north and east into the
Ten Thousand Island zone of the Gulf of Mexico
(Petuch & Roberts 2007). The name Fakahatchee is
derived from the Seminole Indian language which,
roughly translated, means 'muddy creek.' During
the summer rainy season from May to October,
which provides about 42 of the annual 53 inches of
precipitation, most of the Preserve is inundated to
varying depths. This is reversed in winter months


ii


.F.':""' 'I ,?a -"


... P


Fakahatchee
Slrard Presk e-r r
Sli-t- Park -,

.., c ,"l r CoI inl'y


Figure 2. A, County map of Florida showing Collier County and the study site. B, Map of Fakahatchee Strand
Preserve State Park, -n hnii the Tii,; Trail (Route 41) within (c.-.i, 'l-il, the four main collection
sites described in the text are indicated by numerals 1 to 4. Map reprinted with permission of the Florida
Department of Environment Protection, Tallahassee, Florida. Color version available online at ll.p ", 1. ,, l1h1
1ii ,.,.Jii ,iiiii,.[iii! ,,i4"!,,4 ii ii,.- t-2.pdf.


V






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


as ground water recedes and in drier years much of
the Preserve may lack surface water.
Winter temperatures are mild, ranging
normally from 100C to 180C. The mean January
low temperature is slightly below 120 which, in the
strict definition, would place the Preserve slightly
outside the subtropical zone. Occasional cold fronts
may bring mild frosts. Summer months are very
humid with overnight lows rarely dropping below
180C and daytime highs around 320C. The rock
substrate is predominately of biogenetic limestone
(Petuch & Roberts 2007) but outcrops projecting
sufficiently above ground and water levels to support
lichen colonization are rare. Although logged
extensively during the 1940s and 1950s, relatively
mature secondary forests now cover a large portion
of the Preserve. In the north and central part, the
major tree *..... .-. include Taxodium distichum
(bald cypress), Ilex cassine (dahoon), Fraxinus
S i/, h ui (Carolina ash), Roystonia elata (royal
palm), Sabalpalmetto (cabbage palm), Acer rubrum
(red maple) and Quercus i ; 4;'bitn..i (live oak). The
mangrove zone to the south supports Phi-, / h". I
mi,.f,1.: (red mangrove), Avicennia germinans
(black mangrove), L ;:,:J. :.,i i.i racemosa (white
mangrove) and, on higher ground, C ,,. ~_~-.l',
erectus (buttonwood).
Situated between a West Indian-dominated
flora of subtropical Florida to the south and a more
temperate flora to the north, Fakahatchee is known
for its vascular plant diversity by sharing both
floral types (Robertson 1955; Austin et al. 1990).
Encompassing only a little over 85,000 acres
(34,000 hectares), at least 524 vascular plant species
have been recorded from the Preserve (Institute
for Regional Conservation online database: http://
regionalconservation.org/ircs/reports_database.
asp). Outside South Florida, many of these are
found nowhere else in North America and some
are endemic to the park (Avery & Loope 1980;
Austin et al. 1990). By comparison, Everglades
National Park, with its northwest boundary located
just five miles to the southeast, has recorded about
1,200 plant ...;. >-. although it occupies nearly
1.4 million acres (560,000 hectares) (Olmstead &
Loope 1984; Avery & Loope 1996).


Even though the plant flora at the Preserve
has been effectively inventoried, the lichen biota
has not. Due to its climate and geological history,
southern Florida, including the Preserve, possesses
a flora and fauna unlike the rest of the continental
United States (Beard 1938; Robertson 1955;
Olmstead et al. 1981). It is then logical to assume
the lichen biota should also be different. The most
extensive lichen collections from the Preserve
were made in 1992 by Richard Harris and William
Buck and in 1997 by Ralph Common. These forays
produced collections of several hundred species.
A fl-. collections were also taken by Sylvia and
Stephen Sharnoff while accumulating photos for
the book Lichens of K., t,' America coauthored
with Dr. Irwin Brodo in 2001 (Brodo et al. 2001).
Undoubtedly, other unpublished and unrecorded
collections have been made but, if maintained at
all, they appear to be scattered. Currently, no single
database or inventory dedicated to the lichens of
the Preserve is maintained (R. Rau, Park Manager
at Fakahatchee Strand Park Preserve, pers. comm.
2009). The collections previously made by others
plus those gathered during the 2009 Tuckerman
Workshop now document over 400 species and
can be considered representative. This publication
should provide the Preserve with the framework
on which to build a lichen checklist which will,
we hope, be as comprehensive as that of vascular
plants.
COLLECTION SITES
Although small in relation to its giant
neighbors Big Cypress National Preserve and
Everglades National Park, the Preserve contains
many distinct ecological communities including
mangrove forests, prairies of grasses and sedges,
nearly monotypic cypress stands and a variety of
broad leaf forest types. However, due to the limited
collecting time allotted (two full days in addition
to time assigned for laboratory work), only four
sites were selected for sampling. The chosen sites
featured ease of access, a close proximity to one
another and a generous lichen flora.
At Site 1 (Figs. 2B, 3A-B), a lowland forest
is dominated by a unique canopy mix of royal






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


palms and bald cypress. Dahoon and Carolina ash
are common in the subcanopy while the understory
varies from dense to open. In most places at least
moderate sunlight reaches the forest floor. Site 2
features a raised tram road once used by 1,-_..._;_-
companies to extract cut trees (Fig. 3C-D).
Lichens are abundant along the margins of the
road, particularly foliicolous *t .... -.. on cabbage
and saw palm leaves (Sabal ip:i/ii:f i;, Serenoa
.' i,,. ,). Collectors also wandered off into the short
canopy forests on either side of the road. Species of
i; ..! iin ... ; '..//. ;/ and i.,i.. are common
at this site. Site 3 straddles the main road through
the Preserve. It encompasses a nearly monospecific
bald cypress stand on the east side of the road and
an upland hammock (tree island) on the west. The
latter is dominated by live oak, dahoon, cypress
and red maple (Fig. 3E-F). Judging by the large
number of bald cypress stumps and cypress knees,
Site 4 was once a cypress swamp forest (Fig. 3G-
H). Although some '- pr'- has regenerated since
1. '--im days, the site today is dominatedby dahoon,
Carolina ash, red maple, and swamp dogwood
(Cormnusfoemina). The understory was open during
the time of our survey and filtered sunlight reached
the lower strata of the community. During summer
months, site 4 is inundated to various depths but
at the time of collecting ground water was absent
except for a few scattered ponds.
SPECIES IDENTIFICATIONS
Specimens were studied using standard
techniques of light microscopy and thin-layer
chromatography, first during the workshop at
Edison State College in Naples, Florida and
subsequently at the institutions of the participants.
The information was compiled by Rick and
Jean Seavey and Robert Lucking and each
reported taxon was critically revised to ensure
correct identification. In many cases, specimens
were compared with authentic type material,
particularly for the Graphidaceae (including
Thelotremataceae), Porinaceae, Pyrenulaceae, and
Trypetheliaceae. Specimens were studied using a
diverse array of optical t,-,q-ie,-t at Edison State
College and subsequently at the institutions of each
participant. For spot reactions and i-,-,,-,,.-.,-;


staining, we used 10-20% potassium hydroxide
solution (K), commercial bleach containing
6% sodium hypochloride (CHLOROX), para-
phenyldiamine crystals dissolved in 95% ethanol,
and Lugol solution (FLUKA 62650). Standard
thin-layer chromatography (TLC; Amp et al. 1993;
Lumbsch 2002) was applied to a large number
of specimens to confirm spot reactions and to
elucidate more complex chemical patterns. Most
images were taken from freshly collected material
at Edison State College using a NIKON Coolpix
5400 digital camera mounted on one ocular of a
stereomicroscope.
SPECIES RICHNESS
In order to put the number of lichen species
found here into a wider context, we compared the
data to those published for other nature preserves in
the continental United States (Bennett & Wetmore
-'-:.., Spribille et al. 2010) and extracted the
most recent data of the NPLichen database for
national parks and other nature preserves (Bennett
& Wetmore 2005b, Bennett ,^'c^' -; http://www.nbii.
gov/portal/server.pt/community/li chens/1555).
We also used these authors' estimates of relative
0c t 1 1 i -.- -ofthe lichen inventories in these parks
and preserves to account for missing data, knowing
that this can only be an approximation and that
many i..11 .-. require more detailed studies to assess
their lichen diversity accurately. It should be noted
that recent surveys of several p.-l -. considered
90-99% known in the NPLichen database (e.g.
the Santa Monica Mountains and the Great Smoky
Mountains; Knudsen & 2010; Lendemer et al.,
unpubl. data) exhibit undiscovered lichen species
richness that was higher than previously estimated.
However, this applies likely to other parks and
preserves, including Fakahatchee, and therefore
the level of uncertainty is partially balanced out
and the relative comparisons remain valid at this
point.
Assuming that lichen species richness
|l^pe!l-. more or less logarithmically on area and
rather linearly on habitat diversity (McGuiness
1984; Nilsson et al. 1988; Rosenzweig 1995; Ney-
Nifle & Mangel ',,,, McCune & Grace 2, ,,
Lacking et al. 'non,-;. in our analysis we used the







LUCKING ET AL.: Lichens of Fakahatchee Strand Preserve State Park


Figure 3. Habitat photographs of the four collection sites at Fakahatchee Strand Preserve State Park. First row: site 1, showing
royal palm canopy and understory withbald cypress. Second row: site 2, showing side road along old tram with dense vegetation
and aspect of understory with bromeliads. Third row: site 3, showing open bald cypress prairie hammock and dense regrowth.
Fourth row: site 4, showing closed bald cypress forest and aspect of forest understory. Photographs by R. Liicking (A, B, G),
Matthew Nelsen (D, E, H), Frederick Seavey (F), and Jean Seavey (C). Color version available online at http://www.flmnh.ufl.
edu/bulletin/vol49no4/figure3 .pdf.







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. ',, 1,


Table 1. Lichen species richness of selected North American national parks and ..1ib.. pi.. -..' .. -. (after Bennett & Wetmore
2005a, b; Bennett 2006; Spribille et al. 2010; hiup -, -, li l.i I..-. I..l -..- .. i pi ... i aliiii h.. .. i- '1555). SPE = currently
known species richness; COMP = estimate of completeness range; EST = estimated species richness using mean from
estimate of ..i lmpbI.,ii,.. range (mean value; the range would be 5%); SIZE = area cover (excluding water surfaces); ALT
= altitudinal range; VEG = number of vegetation categories present (using simplified classification following Grossman et al.
1 '""... SPE/area = species per area; SPE/log = species per log-area; SPE/veg = species per log-area per vegetation categories.
For further explanation see text. *For these parks, species versus log-area relationships have a steeper slope due to small
area size and hence comparison with larger areas does not I i l, and SPE/area, -.- I ., and SPE/veg calculations are not
given. **Figures updated according to list maintained by park; further 90 species are to be added and hence the lichen species
richness of that park will surpass Kondike (Lendemer, pers. comm. 2010).


PARK NAME SPE COMP


Klondike (Alaska)*
Isle Royale (Michigan)
Great Smoky Mountains (North Carolina)**
Voyageurs (I'hn!! .... :._,.,
Glacier (Montana)
Everglades (Florida)
Acadia (Maine)*
Gates of the Arctic (Alaska)
Fakahatchee S_ unut (Florida)
Y I._., _..... ( ,_m ini ,'
Rocky Mountain (Colorado)
Keweenaw ri iv.. i.II
(,i: mpi.. (W .: ,|l^h _'i..lni
Delaware Water Gap (i i.. Jersey)
Santa Monica Mountains (California)
Chiricahua (Arizona)*
.' ....11. Islands (Wisconsin)
Saint Croix i,' inii. i..1, ,
Saguaro (Arizona)
Big Bend (Texas)
( .11111.I Islands (California)
Pictured Rocks (r' l iI .i..
Denali (Alaska)
Sequoia (California)
Grand Canyon (Arizona)
Theodore Roosevelt (North Dakota)
Redwood (California)
Blue P ,1 I- (North Carolina/Virginia)
Mount Rushmore (South Dakota)*
Grand Teton (Wyoming)
' ,,i, ,d,-i. ,<1 (Virginia)
Hot ,,;..- (Arkansas)*
Bandelier ( i: %, Mexico)*
Costa Rica Las Cruces
Costa Rica La Selva
Costa Rica Tortuguero
Costa Rica Hitoy Cerere
Costa Rica Corcovado


91-99%0
91-99%0
91-99%0
91-99%
91-99%
76-90%
91-99%
91-99%
76-90%
76-90%
91-99%
91-99%
76-90%
91-99%
91-99%
91-99%
91-99%
91-99%
91-99%
91-99%
91-99%
91-99%
51-75%
91-' ..
51-75%
91-' ..
76-'"90,.
51-75%
76-90%
51-75%
76-90%
91-99%
91-99%
91-99%
91-99%
76-90%
76-90%
76-90%


EST SIZE
[1 ,1;]
703 53
645 540
620 2100
525 250
505 3830
540 6000
465 140
465 39000
519 320
505 8500
440 1050
375 8
425 3700
350 265
350 620
340 120
340 160
325 350
315 370
305 3200
285 500
275 290
410 20183
270 1600
405 4900
265 280
300 450
390 370
265 5
350 1250
265 800
225 22
220 130
580 2.7
630 28
630 190
585 200
615 420


ALT [m] VEG SPE/ SPE/ SPE/
area log veg


180-425
265-2025
335-430
960-3190
0-10

85-2525
0-10
1610-3460
2325-4350

0-2430
90-500
0-950

N/A
N/A
665-2640
520-2385
0-550
N/A
70-6195
415-4420
780-2515
590-870
N/A
200-1845


1900-4200 7
180-1235 6


1200-1400
200-400
0-200
200-500
0-200


1.14
0.28
1.99
0.13
0.08

0.01
1.35
0.05
0.40

0.10
1.26
0.54

2.02
0.88
0.81
0.09
0.55
0.91
0.01
0.16
0.05
0.90
0.56
0.66


0.99 0.14
1.37 0.23
1.19 0.17


0.18 0.71 0.10
0.27 0.75 0.13


203.70
21.36
2.75
2.43
1.22


12.75
4.13
2.30
2.11
1.95


12.75
4.13
2.30
2.11
1.95






LUCKING ET AL.: Lichens of Fakahatchee i.i i Preserve State Park


following parameters: total number of species,
-..... .i<, per area, species per log-area, and species
per log-area per habitats. For each preserve, the
total area was corrected to the area covered by
land. To assess the number of principal vegetation
types (suitable for lichen growth) in each preserve,
we applied a simplified classification following
Grossman et al. (1998). In addition to principal
vegetation types, the number of altitudinal belts
was taken into consideration, using a relative
assessment depending on latitude, since the tree
line and hence the potential substrate for epiphytic
lichens decreases with higher latitudes.

RESULTS AND DISCUSSION

Together with previous collections gathered at
Fakahatchee Strand Preserve State Park by R.
Common, W. R. Buck, and R. C. Harris, the
total number of species currently documented
for the Preserve amounts to 432, most of which
were collected during the 2009 Tuckerman
workshop. All species are listed below, together
with representative voucher specimens, and most
of the crustose epiphytic lichens are documented
with color images. The 432 species represent
over 100 genera in 35 families and approximately
2,000 collections in total. Fourteen species new to
science are described below (plus four others in
three separate papers; Nelsen et al. 2010; Lumbsch
et al. 2011; Lucking et al. 2011) and we also report
89 species new to the North American Lichen
Checklist (Esslinger 2010).
Although South Florida lies entirely north of
the Tropic of Capricorn, its floristics are extremely
similar to those of the Caribbean (Tomlinson
1980). Tomlinson also stated that the 120C January
mean temperature isotherm for the state was most
likely the result of oceanic influence and the part
of Florida between the shoreline and this isotherm
would be where the concentration of Caribbean
-...... .i., would be found. With minor exceptions,
this has proved to be the case (Olmstead et al.
1993). Fakahatchee Strand Preserve State Park
narrowly misses being within this zone, but still has
some elements of the Caribbean tree flora such as
Sideroxylon .' JI *, ,. unn1 (mastic),( 'i, 'i--- //in


oliviforme satinleaff) and Randia aculeata
(indigoberry). Conversely, several arboreal species
common at Fakahatchee apparently have reached
their southern limits and become scarce or absent
only a few miles to the south of the site. These
include tFaxinius caroliniana (Carolina ash),
Acer rubrun (red maple), and Quercus i.m! i!. 1.i
(laurel oak). The complete botanical checklist of
the Preserve reveals essentially the same type of
distributional composition and demonstrates that it
is a geographical ecotone between temperate and
tropical biomes.
The 432 lichen species found at Fakahatchee
show a similar mixture of temperate and tropical
taxa, with a distinct bias towards tropical species,
and this certainly accounts for the richness of the
preserve's lichen biota. The majority of the species
recorded here as new to the North American lichen
checklist have tropical affiliations. Some species,
including Pyrenula brunnea, Stirtonia i I,, ../ t.,
and S. dubia, have been known so far only from the
Old World tropics. Others are essentially tropical
but have been collected beyond tropical latitudes
under favorable conditions, such as foliicolous
species (Lendemer & Tripp 2008).
The high percentage of species found to be
new to North America within the preserve's newly
created checklist can in part be attributed to the
i-!'."-1;1 of previous systematic lichen explorations
at this location, as only few collections have
been studied and published before (Harris 1990,
1995). Several lichenologists have collected at
Fakahatchee but there is no record of any lichen
foray ..-itlrt-. the present one. The list below
also includes some taxa that keyed imperfectly or
would not key at all and require further study, most
of them lacking ascomata.
The checklist consists mostly of corticolous
and foliicolous collections with only a few from
lignum or on moss over bark. The lack of saxicolous
specimens is accounted for by the nature of the
geology of the collection site and South Florida
in general. For many million years the area has
been formed by succesive layers of biogenic
marine carbonates, mostly limestone (Randazzo &
Jones 1997; Petuch & Roberts 2007). It has been






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,' 1,


covered by peat and calcareous marls following its
emergence above sea level early in the Pleistocene.
Although limestone outcrops are common, many of
these are inundated nearly half the year by summer
rains. Siliceous rocks are absent from the region.
For the most part, only the higher 3 I),lJr!,-! bc,. ',i
prairies (Orzell & Bridges 2006) contain exposed
calcareous outcrops potentially capable of
supporting a lichen flora. This plant community
was not sampled during this foray.
Although Fakahatchee is a small park, being
22nd in size of all parks with more than 200 lichen
species reported and compared here (Table 1)
and only 10% of the average size of these parks,
it currently ranks 9th in lichen species richness.
By far the largest numbers of species have been
documented for Klondike Gold Rush National
History Park and Isle Royale National Park, with
668 and 613 taxa, respectively (Table 1). Several
other parks have between 400 and 500 species,
including Everglades National Park (Seavey &
Seavey, unpubl. data). In addition to the I..1 .-.
and preserves listed in Table 1, further inventories
are currently under way. Tucker (2010) recently
reported 296 lichen species from Burden Research
Plantation in Louisiana, another subtropical
site that has been monitored for over 25 years.
Hodkinson (2010) and Hodkinson et al. (2009,
2010) found 220 lichen species at Mount Rogers
National Recreation Area.
Excluding parks of less than 200 km2
(because the log-area approximation does not
directly compare between small and large area
sizes due to changes in the slope parameter;
McGuiness 1984; Nilsson et al. 1988; Rosenzweig
1995; Ney-Nifle & Mangel 2000), Fakahatchee
Strand Preserve State Park ranges among the top
three parks for species per log-area and is first in
species per log-area per habitat indices (Table 1).
The latter index surpasses that of much larger parks
with comparable vegetation, such as the nearly
twenty times larger Everglades National Park, and
underlines the carrying capacity of (sub-)tropical
vegetation to support high small-scale diversity,
especially for epiphytic crustose microlichens.


The North American lichen checklist currently
comprises more than 4500 lichen -p.... i,; (Esslinger
2010), which means that Fakahatchee supports
almost 10% of the lichen species richness reported
for the entire North America, although it is only a
fraction of its size (0.00015%). This corresponds
to a 60000-fold increase in species density. This
pl-;,-,-,i-,-,,- of concentrated, high small-scale
species richness comes to its extreme in the tropics,
such as in Costa Rica, where the species per log-area
per habitat snl-,-e are five to ten times larger than
in any -pcI;L.--i l park found in the continental
Unites States, including the (sub-)tropical parts of
Florida (Table 1).

NEW OR OTHERWISE INTERESTING
SPECIES

In the following account, we present taxonomic
novelties found during this and previous lichen
surveys at Fakahatchee. Other new species are
described in separate I,.;.p-' (Nelsen et al. 2010;
Lumbsch et al. 2011; Licking et al. 2011). The taxa
are listed in alphabetical order.

Calopadiafloridana Hodges & Liicking, new
species
Figure 14D-G'
Mycobank #518000

Diagnosis.-Sicut C.,/, puiggarii
apotheciis pruinosis maioribusque differt.
Description.-Thallus corticolous, 1-3 cm
diam., 30-50 ptm thick, centrally continuous but
towardsthe margin with dispersed, irregularpatches;
surface uneven, white to pale gray. Photobiont
chlorococcoid. Apothecia --,I, rounded, 0.6-
1.2 mm diam. and 250-350 pm high; disc plane
to slightly convex, dark gray to brownish gray,
coarsely white-pruinose; margin thin, pale gray to
cream-colored. Excipulum paraplectenchymatous,
50-70 pm broad, colorless. Hypothecium 30-50
pm high, dark brown. A I.-lwil-;., base aeruginous.
Epithecium thin, 5-10 pm high, pale grayish

'Figures 4-58 are available on-line at http://www.
flmnh.ufl.edu/bulletin/vol49no4supplmats.htm






LUCKING ET AL.: Lichens of Fakahatchee t.I i Preserve State Park


brown. Hymenium 120-150 pm high, colorless.
Paraphyses unbranched to slightly branched. Asci
100-120 x 25-30 pm. Ascospores single, ellipsoid,
muriform, 50-80 x 17-25 pm, 3-4 times as long as
broad, colorless. Campylidia sessile, 0.6-1.2 mm
broad, 1-1.5 mm long; lobe well-developed, hood-
shaped, dark gray with paler apex, white-pruinose;
socle not apparent. Conidia filiform with clavate
apex, 5-7-septate, 50-70 x 1.5-2 pm. Secondary
chemistry: no substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Janes Scenic
Drive 6.5 mi NNW of ranger station, west of old
tram, I,. .i /;tli-Saibal hardwood hammock, slough
and strand, on laxodium branch, March 2009,
H 'J'. ., s.n. (F, holotype).
Etymology.-Referring to its discovery in
Florida.
Distribution and Ecology.-This species
is known from two well-developed collections
from Fakahatchee Strand Preserve State Park and
Sumpter County, growing on the smooth part of
the bark of small lixodium branches and partly
on *.. .-.J. areas of the wood bare of periderm, as
well as on hardwood bark.
Remarks.-C,.d,. ',. .i floridana belongs
in a group of species that have single, muriform
ascospores less than 100 pm long. All species so far
known in this group, except the recently described
C. ediae (see below in the -.....- list), have
non-pruinose apothecia which are also distinctly
smaller, usually not exceeding 0.8 mm diam.
Calopadia editae also has pruinose apothecia but
the disc is lighter brown and yellowish pruinose
and the pruina also covers the apothecial margin.
Except for the pruina on the apothecial disc, the
new species comes closest to C. puiggarii, also
present in the park, which shares the dark brown
hypothecium and aeruginous apothecial base and
a gray-brown apothecial disc, but differs in its
smaller, non-pruinose apothecia. Another species
with pruinose apothecia and single-spored asci
is C. i;. i'i-. iimi which has longer ascospores (>
100 pm) and the apothecia, as in C. editae, are
yellow-brown with pale yellow-white pruina
(Lucking 2008). The holotype of C. floridana


bears only mature apothecia, all heavily pruinose,
whereas an additional collection from Sumpter
County (Common 5320K) has younger apothecia
with scarce pruina only, suggesting that the pruina
develops with age.
Additional Specimens Examined.-U.S.A.
Florida. Sumpter Co.: Rt. 471 along Withlacooche
River, at boundary with Polk Co., Apr 1992,
Common 5320K (hb. Common).
Calopadia imshaugii Common & Liicking, new
species
Figure 15A-F
Mycobank #560000
Diagnosis.-Sicut Calopadia fusca sed
apotheciis maioribus pruinosis et ascosporis 1-3 in
ascis differt.
Description.-Thallus corticolous, 0.5-3
cm diam., 30-50 jim thick, centrally continuous
but towards the margin with dispersed, irregular
patches; surface uneven, white to pale gray.
Photobiont chlorococcoid. Apothecia sessile,
rounded, 0.4-0.8 mm diam. and 200-325 pm high;
disc plane to slightly convex, brown, coarsely
white-pruinose; margin conspicuous, pale gray to
cream-colored. IF ,_uilil~t paraplectenchymatous,
50-100 pm broad, colorless. Hypothecium 20-40
pm high, (dark) brown. Apothecial base sordid
brown. Epithecium thin, 5-10 jim high, pale grayish
brown. Hymenium 100-130 pm high, colorless.
Paraphyses unbranched to slightly branched. Asci
90-100 x 20-30 pm. Ascospores (1-)2(-3) per
ascus, ellipsoid, muriform, 50-80 x 15-23 pm,
3-4 times as long as broad, colorless. Campylidia
sessile, 0.5-1 mm broad, 1-1.5 mm long; lobe
well-developed, hood-shaped, dark gray with paler
apex, white-pruinose; socle not apparent. Conidia
filiform with clavate apex, 7-15-septate, 70-90
x 1.5-2 pm. Secondary chemistry: all spot tests
negative.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Scenic Drive
(CR 837)just past bend near gate 14, second growth,
Apr 1997, Common 7322F (MSC, holotype; hb.
Common, i .1 I fK).
Etymology.-We dedicate this new species






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


to the late Henry Imshaug for his contributions to
lichenology.
Distribution and Ecology.-This -.f-ile-. is
known from several collections in Florida, growing
on the bark of small twigs on unidentified trees.
Remarks.-Calopadia .:iiI,,, :ii is unusual
within the genus in having more than one ellipsoid
ascospore per ascus. Most species of the genus
have single-spored asci, and in the fti taxa with
more than one ascospore per ascus, the ascospores
are narrow and oblong rather than ellipsoid, being
70-120 x 8-14 pm (Licking 2008). The ascospores
of the new species have the same shape and
dimensions as those in species with single-spored
asci related to C. ...., Morphologically, the new
species is similar to C. floridana (described above),
but the latter has single-spored asci.
Additional Specimens Examined.-U.S.A.
Florida. Dade Co.: SW 388th St., 1.2 miles east
of Old Dixie Hwy., near Homestead, May 1992,
Common J".."'- ., J '.i ''E (all hb. Common).
(Cha, a platycarpoides (Tuck.) Breuss &
Liicking, new combination
Figure 17D
Mycobank #517995
Basionym.-- 71,i i.. /m, platycarpoides
Tuck., Proc. Am. Acad. Arts Sci. 6:270 (1866);
Phaeotrema platycarpoides (Tuck.) Mull. Arg.,
Flora 69:311. (1886).
Type.-Cuba, Wright 157 (FH!; lectotype,
fide Hale 1981:265).
Remarks.-This species is closely related to
C7'! ".." 7., ,_,,i\..! / with which it co-occurs in the
same microhabitats, mostly on swamp dogwood.
The species was originally separated from C.
platycarpa because of the smaller apothecia and
larger ascospores with more numerous septa.
However, while the two species do differ in
apothecial size, the ascospores are practically
identical, as already noted by Hale (1981). We
found two other distinctive features of C;:...,i/....
Pl, tin .a iJ j. .i, ..' the outer thallus margin is more
or less entire compared to the distinctly lobulate
and recurved margin of C. platycarpa, and the
apothecial margin is UV+ yellow (sometimes


only weakly so), indicating the presence of
lichexanthone. The .t .... -, somewhat resembles
i i, 6ni. i ., porinoides, which also contains stictic
acid, but has much longer, multiseptate, hyaline
ascospores (Rivas Plata et al. 2010).
Additional Specimens Examined: U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Off James Scenic Drive west of Tram
Rd. 6.5 mi NNW of Ranger Station, Taxodium-
Sabal-hardwood hammock, slough, and strand;
March 2009, Breuss 28665 (LI, USF), Licking &
Rivas Plata 26571 (F, USF).
Cjifrlon-i ; midniata Vain. ex Liicking, new
species
Figure 'L F
Mycobank #560001
Diagnosis.-0-)'I, .;!/ !t speciei medulla
cinnabarinadiffert. Thallusecorticatus, virido-albus.
Ascomata desunt. Acido 2-O-methylperlatolico
continens.
Description.-Thallus corticolous, to 10
cm diam., 25-50 pm thick, continuous; surface
uneven, minutely farinose-byssoid, pale green to
almost white, with mottled areas of pure white,
sterile but potentially ascigerous areas; streaks
of medulla frequently exposed, bright orange to
cinnabar-red. Photobiont trentepohlioid. Thallus in
section lacking upper cortex, with irregular algal
layer and areas of concentrated crystalline orange
pigments. Asci and ascospores not observed.
Secondary chemistry: 2-O-methylperlatolic acid
and unknown substance close to gyrophoric in
solvent C, unidentified anthraquinone pigment
detected by TLC (paratype); unpigmented thallus
parts C-, K-, P-, I+ light blue (especially white
areas), pj;i -,,ilel parts K+ dark purple-violet.
Type.-Guyana. Vryheid Plantation, no date
or collector given (TUR-Vainio 28367, holotype).
Etymology.-The epithet refers to the
strongly pigmented parts of the thallus, miniata
meaning cinnabar-red color (not to be confused
with small).
Distribution and Ecology.-This species is
known from the type material from Guyana and
from several collections from Fakahatchee Strand






LUCKING ET AL.: Lichens of Fakahatchee t.i i Preserve State Park


Preserve State Park, where it was found on bark of
Taxodium and hardwood in shaded situations.
Remarks.-The material of this taxon found
in TUR -t._-.-!- that Vainio intended to describe
this species under the generic name Chiodecton,
but never published it. It does not seem to have
been mentioned in the literature. However,
the Florida collections suggest that this is a
widespread, well-defined taxon, even if asci and
ascospores are absent. The general habit, the sterile
white areas dispersed over the thallus showing a
light blue reaction with iodine, and the secondary
chemistry including 2-O-methylperlatolic acid,
support inclusion within C)pi,.''i,,, .,' However,
the pigmented thallus parts are quite unique in the
genus. Thus far, pigments appear to be restricted
to the related genus H1, /" .b! lu (Aptroot et al.
2009), where they usually occur in the hypothallus
and prothallus, as well as the isidioid iiit. n.rx t-..-.
Since the name coined by Vainio was available, we
decided to validate this name rather than using a
new one; therefore, Vainio's material was selected
as holotype. We also specified an epitype from
the Florida material which is then available for
molecular studies.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, off James Scenic Drive 6.5 mi NNW
of ranger station at gate 12 along old tram road,
Taxodium-Sabal hardwood hammock, strand and
..-. on .,v,. i,;,i bark, March 2009, Lay 09-
0063 (hb. Lay), Licking & RA,.. Plata 26678,
26679 (F).
Fissurina aggregatula Common & Liicking,
new species
Figure 23E
Mycobank : .,, '
Diagnosis.-Sicut Fissurina insidiosa sed
lirellis minoribus aggregatibusque differ.
Description.-Thallus corticolous, 1-3 cm
diam., 40-80 rim thick, continuous; surface
uneven to verrucose-bullate, yellowish green to
olive-brown. Photobiont trentepohlioid. Thallus
in section with prosoplectenchymatous upper
cortex, irregular algal layer and scattered clusters


of crystals. Lirellae densely aggregate in small
clusters, straight to curved, unbranched to sparsely
branched, erumpent, with lateral thalline margin,
0.3-0.7 mm long, 0.15-0.2 mm wide, 0.12-0.15
mm high; disc concealed; labia conspicuous, thick,
yellowish-white; thalline margin olive-brown.
Excipulum entire, apically dark brown, 25-50
pm wide; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 5-10 pm high, colorless
to pale yellowish; hymenium 60-80 pm high,
colorless, clear; epithecium granulose, 5-10 jm
high, brown. Paraphyses unbranched, glabrous;
periphysoids not observed; asci fusiform, 60-70
x 10-13 pm. Ascospores 8 per ascus, ellipsoid,
transversely 3-septate, 14-20 x 7-9 pm (including
1-1.5 jam thick wall), 2 times as long as wide,
with comparatively thin septa and angular-rounded
lumina, colorless to pale gray-brown, I-. Secondary
chemistry: no substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, K2 trail
among royal palms, second growth area, Apr 1997,
Common 7356A (MSC, holotype; hb. Common,
isotype).
Etymology.-The epithet refers to the small
clusters of aggregate lirellae.
Distribution and Ecology.-This species
is known from several small collections from
Fakahatchee Strand Preserve State Park, from bark
of branches of unidentified hardwood.
Remarks.-This material is characterized by
erumpent, fissurine lirellae with thicklabiawhich are
clearly I n ii t as well as 3-septate, non-amyloid
ascospores. It comes closest to E insidiosa (Harris
1995; Staiger 2002; Archer 2009), but the lirellae
are much more delicate and form small clusters.
Fissurina radiata is another species with aggregate
lirellae and 3-septate ascospores (see below), but
in that species, the labia are inconspicuous, the
lirellae are .*1j,;,_. and the clusters are much larger
and almost pseudostromatic.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, K2 trail among royal palms, second






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,' 1,


growth area, Apr 1997, Common 7368C-
2 (hb. Common), 7380C-2 (hb. Common).
Fissurina ,,,nnelf'nti, 1, Common & Liicking,
new species
Figure 23F-G
Mycobank #560003
Diagnosis.-Sicut Fissurina i//, i, /i sed
thallo corticato et ascosporis typo 1 i, fli, 1 ; ,- differt.
Thallus corticatus. Lirellae fissurinae, 0.3-0.5 mm
longae, 0.1 mm latae. A. -.,-,-,*.1-,-ae 3-septate, 13-18
x 7-9 jm, I-. Acidi lichenum desunt.
Description.-Thallus corticolous, 0.5-1
cm diam., 30-60 pim thick, continuous;
surface smooth to uneven, pale greenish white.
Photobiont trentepohlioid. Thallus in section with
prosoplectenchymatous upper cortex, irregular
algal layer and clusters of crystals. Lirellae straight
to curved, unbranched to irregularly branched,
ctr ~ilt. fissurine, with thick complete thalline
margin, 0.3-0.5 mm long, 0.1 mm wide, 0.1 mm
high; disc concealed; labia inconspicuous to slightly
_1..* 11w brown along the slit; thalline margin white.
Excipulum entire, apically dark brown, 10-15 pm
wide; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 5-10 pm high, colorless
to pale :I,.h_ ,_.i, hymenium 70-80 pm high,
colorless, clear; epithecium granulose, 5-10 jpm
high, colorless. Paraphyses unbranched, glabrous;
periphysoids not observed; asci fusiform, 60-70
x 12-15 itm. Ascospores 8 per ascus, ellipsoid,
transversely 3-septate, 13-18 x 7-9 ptm (including
1-1.5 pm thick wall), 2 times as long as wide,
with trypethelioid to thin septa and trypethelioid
to anglar-rounded lumina, colorless, I-. Secondary
chemistry: no substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, K2 trail
among royal I..-il-.. second growth area, Apr 1997,
Common 7356N (MSC, holotype; hb. Common,
isotype).
Etymology.-The name is a word play
corresponding to the superficially similar E

Distribution and Ecology.-This species


is known from several rich collections, from
Fakahatchee Strand Preserve State Park and other
parts in Florida. It grows on bark of hardwood.
Remarks.-Fissurina analphabetica belongs
in a complex of species with 3-septate, non-
amyloid ascospores and lacking lichen substances,
centered around E dumastii. Within this complex,
the new species is clearly set apart by its extremely
small and delicate lirellae, which are the smallest
known among all described species. Fissurina
tachygrapha (see below) agrees in the delicate
lirellae with apically dark brown excipulum, as well
as the non-amyloid ascospores with trypethelioid
septa, but has an endoperidermal, ecorticate thallus.
t'issurina illiterata is also superficially similar, but
its thallus is mostly ecorticate and the ascospores
have more rounded lumina and are frequently
weakly amyloid. We also observed conspicuously
thickened paraphyses in that species but not in F
,i~t4 -'h ;l,, 1i- In (
Additional Specimens Examined.--U.S.A.
Florida. Baker Co.: Osceola National Forest
near Ocean Pond, Dec 1975, Common 3673K,
3684H, 3786G (all hb. Common). Collier Co.:
Fakahatchee Strand Preserve State Park, K2 trail
among royal palms, second growth area, Apr 1997,
Common 7356I, 7377E, 7380L (all hb. Common).
Hillsborough Co.: Hillsborough River State
Park, Dec 1990, Common 4667, "-'q'- (all hb.
Common). CR 581, 3.2 mi S of 1-75, Nov 1995,
Common 6601A, '. : : ,, '... !/) (all hb. Common).
Polk Co.: CR 54, 0.1 mi E of CR 557, Apr 1996,
Common 6868P (hb. Common).
Fissurina confusa Common & Liicking, new
species
Figure 24B
Mycobank #560004
Diagnosis.-Sicut Fissurina instabile
sed acido psoromico differt. Thallus corticatus.
Lirellae labiatae, 0.5-1 mm longae, 0.2 mm latae.
Ascosporae muriformes, 25-35 x 12-20 pm, I+
caeruleae. Acido psoromico continens.
Description.-Thallus corticolous, 1-3
cm diam., 50-100 pm thick, continuous; surface
uneven to verrucose-bullate, :;.J1_ ,r.1 green to






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


olive-brown. Photobiont trentepohlioid. Thallus
in section with prosoplectenchymatous upper
cortex, irregular algal layer and scattered clusters
of crystals. Lirellae straight to curved, unbranched
to sparsely branched, erumpent to prominent, with
lateral thalline margin, 0.5-1 mm long, 0.15-
0.2 mm wide, 0.15-0.2 mm high; disc partially
,. .-,, -.. gray-pruinose; labia conspicuous, thick,
white; thalline margin olive-brown. Excipulum
entire, colorless to apically orange-brown, 25-50
ptm wide; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 5-10 pm high, colorless
to pale 9,.1lli.;ll, hymenium 80-100 ptm high,
colorless, clear; epithecium granulose, 5-10 pm
high, brown. Paraphyses unbranched, glabrous;
periphysoids not observed; asci fusiform, 80-90
x 25-35 itm. Ascospores 8 per ascus, ellipsoid,
muriform with 5-7 x 1-3 septa, 25-35 x 12-20
[tm (including 1-2 jtm thick wall), 2 times as
long as wide, with comparatively thin septa and
angular-rounded lumina, colorless, I+ violet-blue.
Secondary chemistry: psoromic acid (major),
subpsoromic acid (minor), 2'-O-demethylpsoromic
acid (major), medulla P+ yellow.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, K2 trail
among royal palms, second growth area, Apr 1997,
Common 7380D (MSC, holotype; hb. Common,
isotype).
Etymology.-The epithet refers to the fact
that this *- .... i<-, was previously confused with E

Distribution and Ecology.-Known from
several collections from Fakahatchee Strand
Preserve State Park and also reported from Brazil
by Staiger (2002; under T. instabilis). The species
is usually found on branches of hardwood.
Remarks.-The Florida material agrees well
with the description of Fissurina instabilis given
by Staiger (2002), including the verrucose-bullate
thallus and the erumpent to prominent lirellae
with slightly '_.'"'"_' disc and thick, white labia.
Staiger (2002) accepts forms with and without
psoromic acid in the same species, but we consider
this premature until molecular data clarify the


situation. In other genera of Graphidaceae, such
as Ocellularia s.lat., psoromic acid is accepted as
a species-diagnostic character. Since the type of
F instabilis has no substances, we introduce here
the name Fissurina .i:/r.,, for the psoromic-
acid containing specimens. All Florida material
contains psoromic acid. Fissurina presently
contains four species with muriform (and amyloid)
ascospores and psoromic acid chemistry. Among
these, Fissurina ;:]i,;.~7ji. (Nyl.) Staiger is set
apart by the very small, almost globose ascospores
(about 10 x 10 jim), whereas Fissurina streimannii
(A. W. Archer) A. W. Archer has more elongate
ascospores (about 18 x 10 ptm); both have closed,
labiate lirellae with white upper surface. The other
two -'....<.-. have comparatively large ascospores
(about 30 x 15 pm) and differ from each other
morphologically: Fissurina columbina (see below)
has erumpent, fissurine lirellae with inconspicuous
labia, whereas Fissurina psoromica (A. W. Archer)
A. W. Archer has labiate lirellae with concealed
disc which are brown-black along the slit due to
a weakly carbonized excipulum; in addition, that
species differs in having 2-methoxy-psoromic
acid, a substance that can be distinguished from
psoromic acid only by HPLC (Archer 2001). The
new species, F . .:-ts, ., also has labiate lirellae but
with partially exposed disc and white labia, thus
clearly different from both E' columbina and F
psoromica.
Additional Specimens Examined.--U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, K2 trail among royal palms, second
growth area, Apr 1997, Common 7356A (hb.
Common), 7356F (hb. Common), 7368G (hb.
Common).
Fissurina inspersa Common & Liicking, new
species
Figure 25B
Mycobank #560005
Diagnosis.-SicutFissurina i.., .- l 'l, .. i.
hymenio insperso et acido stictico differt. Thallus
pro parte ecorticatus. Lirellae fissurinae, 0.5-1 mm
longae, 0.1 mm latae. Ascosporae 3-septate, 12-18
x 6-9 jim, I-. Acido stictico continens.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


Description.-Thallus corticolous, 0.5-1
cm diam., 30-60 jpm thick, continuous; surface
smooth to uneven, yellowish to pale greenish
white. Photobiont trentepohlioid. Thallus in
section with partly prosoplectenchymatous, partly
irregular and loose upper cortex, irregular algal
layer and scattered clusters of crystals. Lirellae
straight to curved, unbranched, immersed,
fissurine, with thick complete thalline margin,
0.5-1 mm long, 0.1 mm wide, 0.1 mm high;
disc partially exposed, dark gray-pruinose; labia
inconspicuous; thalline margin yellowish white.
Excipulum entire, brown to apically dark brown,
10-15 jtm wide; laterally covered by corticate
algiferous thallus including clusters of crystals;
hypothecium prosoplectenchymatous, 5-10 [im
high, colorless to pale yellowish; hymenium
50-60 ptm high, colorless, strongly and densely
inspersed (inspersion persistent in K); epithecium
granulose, 5-10 pm high, brown. Paraphyses
unbranched, glabrous; periphysoids not observed;
asci fusiform, 50-60 x 10-12 tm. Ascospores 8
per ascus, ellipsoid, transversely 3-septate, 12-
18 x 6-9 ptm (including 1-1.5 jim thick -...ill), 2
times as long as wide, with trypethelioid septa and
lumina, colorless, I-. Secondary chemistry: stictic
acid (major), constictic acid (trace), thallus section
with K+ yellow <. ffl u.. under the microscope.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Scenic Drive
(CR 837)j t f.. t bend near gate 14, second growth,
Apr 1997, Common 7323C (MSC, holotype; hb.
Common, isotype).
Etymology.-The name refers to the strongly
inspersed hymenium, an unusual character in
Fissurina (Staiger 2002).
Distribution and Ecology.-This species
is known from several rich collections, all from
Fakahatchee Strand Preserve State Park, on
hardwood.
Remarks.--Fissurina inspersa is a typical
Fissurina with an unusual character: a strongly
inspersed hymenium. The inspersion resembles
that of Graphis cinerea and allies (Lacking 2009),
in that it makes the paraphyses, asci, and ascospores
barely discernible, but contrary to the latter, in


tFissurina inspersa it does not dissolve in K. Thus far,
no species of Fissurina with inspersed hymenium
appears to be known (Staiger 2002). Apart from
the inspersion and the darkened ..,_ipukln1
F'issurina dumastoides Fink comes closest, as it
shares the fissurine lirellae of type I according to
Staiger (2002), has the same ascospore type, and
also produces stictic acid. Fissurina iL,. I// ,i ,, */..'
(see below) looks superficially similar and also
has a weakly carbonized l.._ipur.li and the same
ascospore type, but lacks inspersion and its thallus
is endoperidermal and completely ecorticate.
Additional Specimens Examined.--U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Scenic Drive (CR 837) just past bend
near gate 14, second growth, Apr 1997, Common
7276F (hb. Common); ibid., K2 trail among royal
palms, second growth area, Apr 1997, Common
73560 (hb. Common), 7368J (hb. Common).

Fissurina mexicana (Zahlbr.) Liicking & Rivas
Plata, new combination
Figure 25C
Mycobank #517999
Basionym. -Enlerodiclyon mexicanum
Zahlbr., Ann. Mycol. 19:233 (1921); 0Cii_"1,,t
mexicana (Zahlbr.) M. Wirth & Hale, Contr. US
Natl. Herb. 36:82 (1963).
Type.-Mexico, Pringle 20 (W).
Remarks.-This species is usually identified
with Fissurina i,,, li Nyl. (Harris 1995).
Staiger i(-"'.') noted that FE. i, ,1, ,..! I., was
described as sterile, and indeed, no ascospores are
present in the type (indicated on Mason Hale's
Index Cards at US). The application of this name
is therefore uncertain. There are two other species
with similar thallus and lirellae morphology:
GC.'/; '-. -_,_-',.., f. intercludens Nyl. and GI.C'/, ":,.
mexicana (Zahlbr.) Wirth & Hale. Ascospores are
present in both types, being 20-25 pm long in
intercludens and 25-40 pm long in mexicana. The
first was listed as a synonym ofFissurina egena by
Staiger, but differs in the pseudostromatic lirellae
and therefore is recombined inFissurina: Fissurina
intercludens (Nyl.) Liicking & Rivas Plata, new
combination and status [Mycobank #517996;






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


Basionym.-Graphis egena f. intercludens Nyl.,
Bull. Soc. Linn. Normandie, Ser. 2, 2:121 (1868).
Type.-New Caledonia, V1iellard s.n. (H-NYL
7455!; lectotype, fide Staiger 2002:128)]. In the
material from Fakahatchee, ascospores are mostly
30-45 jim long and fit Graphina mexicana. Since
the type of F !r,: .,l,. l., is from SE Asia
(Bengal), we consider it highly unlikely that F
leuconephela is conspecific with G. mexicana.
Harris (1995) gave the ascospore size for Fissurina
leuconephela as 35-78 x 18-36 pm. We have not
seen ascospores of such a size and they seem to
be highly unusual for a Fissurina; probably there
is a further species belonging to a different genus
involved. The collection Common 7323H bears
pycnidia with bacillar conidia about 5 1 Im in size.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Scenic Drive (CR 837) just past bend
near gate 14, second growth, Apr 1997, Common
7323H). Along James Scenic Drive, 3.2 mi NNW
of Ranger Station; 7i.,.l ,,,nii h.-iiwood strand,
swamp and slough, Mar 2009, Ll ..l w; & Rivas
t 'I. 1 26513*, 26514, 26515 (all F).
Fissurina pseudostromatica Liicking & Rivas
Plata, new species
Figure 25D
Mycobank -,00_I.,
Diagnosis.-Sicut F'issurina,..:-:, .". ,' ,,,. < sed
lirellis immersis albis et ascosporis angustioribus
differt. Thallus corticatus. Lirellae fissurinae, 0.3-
0.7 mm longae, 0.15-0.2 mm latae. Ascosporae
3-septate, 14-20 x 7-9 pm, I-. Acidi lichenum
desunt.
Description.-Thallus corticolous, 1-3 cm
diam., 40-60 pm thick, continuous; surface smooth
to uneven, olive-green. Photobiont trentepohlioid.
Thallus in section with prosoplectenchymatous
upper cortex, irregular algal layer and scattered
clusters of crystals. Lirellae densely aggregate
in pseudostromatic, white clusters, straight to
curved, unbranched, immersed, with thin thalline
margin, 0.1-0.3 mm 1.!_,-.. 0.1 mm wide, 0.1 mm
high; disc concealed to slightly gaping; labia
inconspicuous, white; thalline margin yellow-


white. Excipulum entire, orange-brown, 15-20
jim wide; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 5-10 jim high, colorless
to pale yellowish; hymenium 60-80 pm high,
colorless, clear; epithecium granulose, 5-10 Lim
high, orange-brown. Paraphyses unbranched,
glabrous; periphysoids present, glabrous; asci
fusiform, 60-80 x 10-15 pm. Ascospores 8 per
ascus, ellipsoid, transversely 3-septate, 15-20 x
5-7 im (including 1 im thl-,,- -ill), 3 times as long
as wide, with comparatively thin septa and angular-
rounded lumina, colorless to pale gray-brown, I-.
Secondary chemistry: no substances detected by
TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Janes Scenic
Drive 6.5 mi NNW of ranger station, west of old
tram, i', Jin,,,i Sabal hardwood hammock, slough
and strand, March 2009, ZI,.ii & Rivas Plata
26512 (F, holotype).
Etymology.-The epithet refers to the
pseudostromatic clusters of aggregate lirellae.
Distribution and ecology.-This species is
known from a single collection from Fakahatchee
Strand Preserve State Park, found on branches of
hardwood.
Remarks.-This material was first considered
to represent Fissurina mexicana, another species
with pseudostromatic clusters oflirellae (see above).
However, 1F mexicana has muriform ascospores and
its thallus is ligher, with a less conspicuous contrast
between thallus and pseudostromatic clusters.
Fissurina pseudostromatica appears to be the only
species known in the genus with pseudostromatic
clusters and transversely septate ascospores.
Fissurina tuckermaniana Common & Liicking,
new species
Figure 26B
Mycobank #560007
Diagnosis.-Sicut Fissurina comparile sed
,1 -,-:i.f- ; ; m aioribus et p.-.-, ,l -, 1-.;; *. spinulosis
differt. Thallus corticatus. Lirellae fissurinae, 1-3
mm longae, 0.25 mm latae. Ascosporae muriformes,
15-25 x 6-8 im, I-. Acidi lichenum desunt.






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


Description.-Thallus corticolous, 1-2 cm
diam., 30-50 jtm thick, continuous; surface
smooth, yellowish-brown to olive-brown. Photo-
biont trentepohlioid. Thallus in section with
prosoplectenchymatous upper cortex, irregular
algal layer and scattered clusters of crystals.
Lirellae straight to curved, unbranched to sparsely
branched, immersed to erumpent, fissurine, with
complete thalline margin, 1-3 mm long, 0.3-0.5
mm wide, 0.15-0.2 mm high; disc concealed;
labia conspicuous but not thickened, grayish black
to brown-black below corticate thalline margin;
thalline margin olive-brown. Excipulum entire,
brown-black, 25-50 ptm wide; covered by corticate
algiferous thallus including clusters of crystals;
hypothecium prosoplectenchymatous, 5-10 [im
high, colorless to pale yellowish; hymenium
80-100 pm high, colorless, clear; epithecium
granulose, 5-10 pm high, gray-brown. Paraphyses
unbranched, apically spinulose; periphysoids
present, apically spinulose; asci fusiform, 80-90
x 20-25 tm. Ascospores 2-4 per ascus, ellipsoid,
transversely 3-septate, 25-40 x 11-14 jpm
(including 1-1.5 [im thick ..ill), 2-3 times as long
as wide, with trypethelioid septa and lumina,
colorless, I+ violet-blue. Secondary chemistry: no
substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Scenic Drive
(CR 837) just past bend near gate 14, second growth,
Apr 1997, Common 7323D) (MSC, holotype).
Etymology.-We honor Edward Tuckerman
for his contributions to North American lichenology
with this new species.
Distribution and Ecology.-This species
is known from two rich collections on hardwood
from Fakahatchee Strand Preserve State Park.
Remarks.-Fissurina luckermaniana
externally resembles Fissurina comparilis (Nyl.)
Nyl. and Fissurina ,q; ,nne..r. Staiger
(Staiger 2002) in the corticate, smooth, olive-
brown thallus and the fissurine, carbonized lirellae.
Anatomically, however, the species is quite
unique in having unusually large, albeit 3-septate
ascospores in numbers of 2-4 per ascus and apically
spinulose paraphyses and periphysoids. Spinulose


paraphyses do occur in a few other species in
Fissurina (Staiger 2002), but not in combination
with carbonized lirellae (except for the new
species Fissurina .iifl.' ,wiw,.. ,;id ,i,.. (formally
described in Lumbsch et al. 2011). The ascospores
are the largest 3-septate ascospores thus far known
in Fissurina. Staiger (2002) gives Fissurina
lia I_ ~' ..oq .LI' (Nyl.) Staiger as species similar to F
i,,,'. .ij but with larger ascospores; however,
F it .- ;..i'l/;i clearly differs in the largely
endoperidermal, pale yellowish-gray thallus and
the only weakly and apically carbonized lirellae; it
also lacks spinulose paraphyses and periphysoids
and is thus is quite distinct from E tuckermaniana.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Scenic Drive (CR 837) just past bend
near gate 14, second growth, Apr 1997, Common
7266 (hb. Common).
Fissurina varieseptata Common & Liicking,
new species
Figure 26C
Mycobank #560008
Diagnosis.-Sicut Fissurina illiterate sed
ascospores 7-septatis differt. Thallus pro part
ecorticatus. Lirellae fissurinae, 0.5-1 mm longae,
0.1 mm latae. Ascosporae 5-7-septate, 12-18 x
4-5 pm, I-. Acidi lichenum desunt.
Description.-Thallus corticolous, 1-5
cm diam., 40-70 pm thick, continuous; surface
smooth to uneven, .11l,.;*h white. Photobiont
trentepohlioid. Thallus in section with irregular
and loose upper cortex, irregular algal layer and
clusters of crystals. Lirellae straight to curved,
unbranched to sparsely branched, immersed,
fissurine, with complete thalline margin, 0.5-1 mm
long, 0.1 mm wide, 0.1 mm high; disc concealed;
labia inconspicuous; thalline margin yellowish
white but yellow-brown along the slit. E ..-i,_uluki
entire, colorless to apically orange-brown, 15-25
jtm wide; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 5-10 ipm high, colorless
to pale yellowish; hymenium 60-80 pm high,
colorless, clear; epithecium granulose, 5-10 jim






LUCKING ET AL.: Lichens of Fakahatchee t.I i Preserve State Park


high, gray-brown. Paraphyses unbranched,
glabrous; periphysoids not observed; asci fusiform,
60-70 x 10-15 pm. Ascospores 8 per ascus,
ellipsoid, transversely (3-)5-7-septate, 12-18 x 4-5
pm (including 0.5-1 pm thick wall), 3-4 times as
long as wide, with trypethelioid septa and lumina,
colorless, I-. Secondary chemistry: no substances
detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Big Cypress
Bend, old growth Cypress swamp, Apr 1997,
Common 7413A (MSC, holotype).
Etymology.-The name refers to the
transversely multiseptate ascospores.
Distribution and ecology.-This species
is known from a single collection, growing on
hardwood, from Fakahatchee Strand Preserve State
Park.
Remarks.--Fissurina . i ti is
superficially similar to Fissurina illiterata and
perhaps closely related to that species. It differs,
however, in the mostly 5-7-septate ascospores, a
highly unusual trait in the genus, where the species
known to date have either 3-septate or (sub-)
muriform ascospores (Staiger 2002). In spite of
careful search, we have not found any indication in
the literature or studied type specimens that -r ,--_. t
a previous description of a multiseptate species
that would belong in Fissurina. On the other hand,
most species with muriform ascospores have 5-7
transverse primary septa.
Graphis appendiculata Common & Liicking,
new species
Figure 27D-E
Mycobank #560009
Diagnosis.-Sicut Graphide olivacea sed
ascosporis minoribus et thallo albo-cinereo differt.
Description.-Thallus corticolous, 1-3 cm
diam., 50-100 jm thick, continuous; surface smooth,
white to pale gray. Photobiont trentepohlioid.
Thallus in section with prosoplectenchymatous
upper cortex, irregular algal layer and large clusters
of crystals. Lirellae flexuose, irregularly branched
but often in radiating lines, immersed to erumpent,
with lateral to apically thin complete thalline margin,


3-7 mm long, 0.1-0.2 mm wide (abruptly thinner
at the ends), 0.15-0.2 mm high; disc concealed;
labia finely but distinctly striate, black or dark gray
(when covered by thin thallin margin); thalline
margin white to pale gray. Excipulum crenulate,
apically carbonized and laterally and basally
orange-brown, 30-50 pm wide and basally to 25
pm high; laterally covered by corticate algiferous
thallus including clusters of crystals; hypothecium
prosoplectenchymatous, 10-15 pm high, colorless
to pale yellowish; hymenium 100-150 pm high,
colorless, clear; epithecium granulose, 5-15 pm
high, gray-brown. Paraphyses unbranched; asci
fusiform, 80-120 x 30-40 [im. Ascospores 8 per
ascus, oblong, transversely 9-13-septate, 40-60 x
9-13 ipm, 4-6 times as long as wide, colorless, I+
violet-blue. Secondary chemistry: no substances
detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Scenic Drive
(CR 837)just past bend near gate 14, second growth,
Apr 1997, Common 7313A (MSC, holotype; hb.
Common, isotype).
Etymology.-The name refers to the peculiar
shape of the lirellae, which become abruptly narrow
and with entire labia towards the ends and hence
appear appendiculate.
Distribution and Ecology.-Surprisingly this
new species is one of the most commonly collected
graphids at Fakahatchee but not yet known from
outside the area, making it a possible local endemic.
It mostly grows on small branches of hardwoods in
partially exposed microhabitats.
Remarks.-With a number of over 300
accepted taxa and well over 1000 described names,
describing new .p .... .. in the genus Graphis is
problematic. Nevertheless, in spite of careful
search, we have not found a name for this taxon,
which is represented by abundant material. With
the immersed-erumpent lirellae with striate labia
and apically carbonized excipulum, together
with the clear hymenium, transversely septate
ascospores, and no chemistry, it comes closest
to Graphics subalbostriata Lu.l.in _. Graphics
olivacea Redinger, Graphis / ,l ,'/, ~i., Mill. Arg.,
and Graphis proserpens (Lul.iit et al. 2009).






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,' 1,


However, both G. subalbostriata and G. olivacea
have ascospores over 80 lim long, whereas in G.
,,/i ;.. i, and G. proserpens, they are 20-40 pm
long. The often present thin cortex covering the
labia apically is also found in G. ,h i. ..,, but
besides the larger ascospores, that species also
differs by its dark, olive-gray thallus. Graphics
..o '-i.n .ij.; IJ..II can be considered a further species
in the difficult G. proservy i- t ,l :i. '/,L, complex.
Graphics ,.y,'.. /../il./i, is the most abundant
Graphis at Fakahatchee. The collections made
during the Tuckerman workshop in March 2009
are almost without exception -ril ii, whereas those
made by R. Common in Apr 1997 are abundantly
fertile, suggesting notjust a seasonality in ascospore
production but periodicity that could span more
than a year's cycle. The species does not key out
in Harris (1995); the most similar species there is
Graphis /,1', .L -,:/.. Nyl., which has prominent to
sessile lirellae and smaller ascospores, but it too is
listed by Harris (1995) as frequently sterile.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Scenic Drive (CR 837) just past bend
near gate 14, second growth, Apr 1997, Common
7318K (hb. Common); ibid., K2 trail among royal
palms, second growth area, Apr 1997, Common
7368L, 73801 (all hb. Common). U.S.A. Florida.
Collier Co.: Fakahatchee Strand Preserve State
Park, off James Scenic Drive 6.5 mi NNW of ranger
station at gate 12 along old tram road, 7axodium-
Sabal hardwood hammock, strand and swamp, on
7:,\,.. i,,, bark, March 2009, Lay 09-0068 (hb.
Lay), l 1' ur: & Rivas Plata 26657, 26659, 26660,
26668 (all F, USF).
Phaeographis delicatula Common & Liicking,
new species
Figure 44D
Mycobank : ;-.-,,10
Diagnosis.- Sicut ,1.'. ,, i, ,i,;;. brasili-
ense sed acidum sticticum continent differt.
Description.-Thallus corticolous, 1-3
cm diam., 50-100 jtm thick, continuous; surface
uneven-verrucose, pale yellow-gray to olive-
yellow. Photobiont ,i)ftL.pld i section with prosoplectenchymatous upper cortex,


irregular algal layer and large clusters of crystals.
Lirellae flexuose, aggregate in dense clusters but
not pseudostromatic, stellately branched, erumpent,
with indistinct labia and thin thalline margin, 1-3
mm long (but sometimes secondarily divided into
shorter segments), 0.1 mm wide, 0.12-0.15 mm
high; disc exposed, chocolate-brown, sometimes
with thin gray pruina; labia indistinct, dark brown;
thalline margin yellow-white. Excipulum divergent,
dark brown, 10-20 tmm wide; laterally covered
by corticate algiferous thallus with clusters of
crystals between individual lirellae; hypothecium
prosoplectenchymatous, 10-15 ptm high, colorless
to pale yellowish; hymenium 50-60 pm high,
colorless, clear; epithecium granulose, 5-10 pm
high, gray-brown. Paraphyses unbranched; asci
fusiform to clavate, 45-55 x 10-12 pm. Ascospores
8 per ascus, oblong, transversely 3 i-pt.atr, 13-18
x 5-7 pm, 2-3 times as long as wide, colorless, I+
vine-red to red-purple. Secondary chemistry: stictic
acid (major), constictic acid (trace), thallus section
with K+ yellow efflux under the microscope.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, K2 trail
among royal palms, second growth area, Apr 1997,
Common 7367C (MSC, holotype; hb. Common,
isotype).
Etymology.-The name refers to the delicate
lirellae.
Distribution and Ecology.-Known from
three collections from Fakahatchee Strand Preserve
State Park, all on branches of hardwood.
Remarks.-The clear hymenium, aggregate-
stellate lirellae, and small ascospores indicate
this new species to belong in the /'.i,. ..' ',v,,,/
intricans .,.i-...'.,i: (Staiger 2,"'C-). Most species
in this aggregate have norstictic acid; thus far only
one species, itii..1. /.. i,,. -i, flavescens, is known
with stictic acid (Dal Fomo & Eliasaro 2010). The
latter has more robust, distinctly pruinose lirellae
and larger, n- i.1l, 5-septate ascospores.
Additional Specimens Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, K2 trail among royal palms, second
growth area, Apr 1997, Common 7355J (hb.
Common), 7381D (hb. Common).






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


Tapellariafloridensis Common & Liicking, new
species
Figure 56C
Mycobank :: _.-011
Diagnosis.-Sicut 7q/l'. oI.n r malmei sed
apotheciis marginibus pruinosis et ascosporis
minoribus differt.
Description.-Thallus corticolous, 1-2 cm
diam., 30-50 pm thick, continuous; surface uneven,
white to pale gray. Photobiont chlorococcoid.
Apothecia sessile, rounded, 0.3-0.7 mm diam. and
170-270 pim high; disc at first concave, eventually
plane to slightly convex in old apothecia, black;
marginthick, prominent, evanescentin old apothecia,
thickly gray-pruinose especially in younger
apothecia. Excipulum paraplectenchymatous, 30-
50 pm broad, dark purplish-brown. Hypothecium
30-60 pm high, dark purplish brown, K+ purplish.
Apothecial base dark purplish brown. Epithecium
10-15 pim high, blackish brown. Hymenium 100-
120 pm high, colorless or light purplish in upper
parts. Paraphyses branched and anastomosing.
Asci 90-110 x 18-25 pm. Ascospores 4-8 per
ascus, ellipsoid, muriform with 3-5 transverse
and 0-2 longitudinal septa per segment, 20-25 x
9-12 pim, 2-2.5 times as long as broad, colorless.
Campylidia sessile, 0.4-0.5 mm broad, 0.6-0.8
mm long; lobe well-developed, hood-shaped,
black; base not thickened. Conidia filiform, curved,
7-13-septate, 80-90 x 2-2.5 pm. Secondary
chemistry: no substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Pi. i- State Park, Scenic Drive
(CR 837)just past bend near gate 14, second growth,
Apr 1997, Common 7 -." i (MSC, holotype, USF,
hb. Common, isotypes).
Etymology.-Refers to the state of Florida.
Distribution and Ecology.-This species is
known from two, well-developed collections from
Fakahatchee Strand Preserve State Park, growing
on branches and trunks of hardwood.
Remarks.--1"/, 4/i. i, i, t is a chiefly foliicolous
genus characterized by lecideine apothecia,
Byssoloma-type asci, anastomosing, net-like
paraphyses, thin-walled, transversely septate
to muriform ascospores, and campylidioid


conidiomata producing filiform, multiseptate
conidia (Lu .i.g 2008). The .t..... are
distinguished chiefly by their ascospore septation
and size and the absence or presence of pruina on
the apothecial margin. Thus far, only one species,
Tapellaria malmei, was known to produce small,
muriform ascospores in numbers of 4-8 per ascus
(Lucking 2008). T.. h./iii,i floridensis species
agrees with i malmei in ascospore type, but
the young apothecia are epruinose in T malmei
and thickly gray-pruinose in the new species; in
addition, the ascospores are larger in T malmei
(25-35 x 14-20 pm).
Additional Specimen examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Scenic Drive (CR 837) just past bend
near gate 14, second growth, Apr 1997, Common
7315D (hb. Common).

Tilidrlia granulosa Liicking & Rivas Plata,
new species
Figure 56D
Mycobank #560012

Diagnosis.-Sicut 7.' /i., i., malmei sed
thallo granuloso differ.
Description.-Thallus corticolous, 2-5
cm diam., 30-50 pm thick, continuous; surface
finely and densely granulose-isidiate, pale green;
granules ('isidia') formed by agglomerated algal
cells wrapped in fungal hyphae. Photobiont
chlorococcoid. Apothecia sessile, rounded to
irregular in outline, 0.5-1 mm diam. and 180-
270 pm high; disc plane becoming convex, black
to brownish black; margin thin but distinct,
not prominent, persistent, black. E ..,_iulukM
paraplectenchymatous, 30-50 pm broad, gray-
brown. Hypothecium 50-100 pm high, dark
purplish brown, K+ purplish. Apothecial base
dark purplish brown. Epithecium 10-20 jim high,
blackish brown. Hymenium 100-130 pm high,
colorless or light purplish in upper parts. Paraphyses
branched and anastomosing. Asci 90-120 x 18-28
pm. Ascospores 4-8 per ascus, ellipsoid, muriform
with 3-5 transverse and 1-2 longitudinal septa per
segment, 20-25 x 10-15 pm, 1.8-2 times as long
as broad, colorless. Campylidia sessile, 0.4-0.6






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,'I 1


mm broad, 0.7-1 mm long; lobe well-developed,
hood-shaped, dark gray to black with white-gray
base; base not thickened. Conidia filiform, curved,
5-7-septate, 40-50 x 2 [tm. Secondary chemistry:
no substances detected by TLC.
Type.-U.S.A. Florida. Collier Co.: Faka-
hatchee Strand Preserve State Park, Janes Scenic
Drive 1.8 mi NNW of ranger station, Ti ..in intt-
Sabal hardwood prairie hammock, March 2009,
Liicking & REj;, I'/.i0 26810 (F, holotype; USF,
isotype).
Etymology.-The epithet refers to the finely
and densely granulose thallus, an unusual feature
in the genus Tipy. i11 e../i
Distribution and Ecology.-This -p&f-i-; is
known from two well-developed collections from
Fakahatchee Strand Preserve State Park, growing
on ,% oI.. ../; / bark.
Remarks.-This is a second new species
related to Tapellaria malmei, which is also found
at the type locality. Like the proceeding species,
iT'i../ l i,..i granulosa shares with T. malmei the
small, muriform, broadly ellipsoid ascospores
occurring in numbers of 4-8 per ascus. However,
the finely and densely granulose-isidiate thallus
differs from all T.', ,..1l /I.. species known to date. It
is unlikely that these granules serve as isidia; rather
they should be interpreted as increasing the thallus
surface. A similar interspecific variation in thallus
surface structure is known from the related genus
Lasioloma (Lucking & Serusiaux 2001).
Additional Specimen Examined.-U.S.A.
Florida. Collier Co.: Fakahatchee Strand Preserve
State Park, Janes Scenic Drive 1.8 mi NNW of
ranger station, 7L,'\., Jin,,, Sabal hardwood prairie
hammock, March 2009, Lay 09-0013 (hb. Lay),
Licking & Rivas Plata 26697 (F), Mercado-Diaz
394 (UPR).

ANNOTATED CHECKLIST OF LICHEN
SPECIES FROM FAKAHATCHEE STRAND
PRESERVE STATE PARK

In the following list, genera and species are
listed in alphabetical order. Brief taxonomic and
nomenclatural annotations are given for selected
taxa. Selected and representative specimens are


cited for each taxon in parentheses after the species
name. Herbaria are not indicated in the list but
are as follows: DUKE (B. P. Hodkinson), F (R.
Lacking & E. Rivas Plata, M. P. Nelsen), FTU (W.
Safranek), ILLS (L. Crane), LI (0. Breuss), MSC
(R. Common, types and new records for United
States), NY (S. Q. Beeching p.p., W. R. Buck, R.
C. Harris, J. C. Lendemer), OAC (T. McMullin),
UPR (J. A. Mercado-Diaz), and the private herbaria
of S. Q. Beeching, R. Common, M. Hodges, E.
Lay, W. B. Sanders, H. P. Schaefer, and R. & J.
Seavey. Where the material allowed, selected
duplicate specimens are deposited at the University
of Southern Florida in Tampa (USF); these are
marked with an asterisk (*) after the collection
number. Most crustose species are illustrated
by color photographs in a series of plates in the
supplemental online appendix (Figs 4-58; http://
www.flmnh.ufl.edu/bulletin/vol49no4supplmats.
htm); for a few taxa for which the Fakahatchee
material did not allow to produce good images, we
replaced these by images taken from specimens
from other regions. For selected genera that are
either not treated in detail in Harris (1990, 1995)
or more recent publications (Ekman 1996; Staiger
& Kalb 1999; Marbach 2000; Lucking et al. 2007;
Lendemer et al. 2009; Seavey 2009; Seavey &
Seavey 2009; Tripp et al. 2010) or for which many
new records are cited, we provide brief keytables
that summarize the characters of each -p1... i<.-. and
allow their identification and differentiation. In
these keytables, to ... <.., are arranged in taxonomic
order by column characters from left to right; to
key out a species, one would therefore start with
the first column and then use subsequent columns
to narrow down the identification. Species new to
the North American lichen checklist are marked
with an asterisk (*) before the species name; some
of these have already been known to occur in North
America but have either not been reported in the
literature or not yet added to the i- hi,- tI -r Species
newly reported to the New World are preceded by
two asterisks (**) and new species (described above
or in separate papers) with three asterisks(***).
Acanthothecis peplophora (M. Wirth & Hale)
E. A. Tripp & Lendemer-Figure 4A-B






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


(Beeching 7 's-, Common 7323G, Crane ILLS
.;i031, Lay 09-0038, Lendemer 15673, Ln.. 1; -
& Rivas Plata 26500*, 26503*, Mercado-
Diaz 4-'1,- Seavey & Seavey 11007*). Material
from Florida was sequenced and clusters with
Graphina aff. peplophora from Cuba (Staiger
et al. 2006; Rivas Plata et al., unpubl. data).
Thus far, no other species of A...., l1i di...i..
have been sequenced, but based on the available
data, the genus is distinct from other lirellate
Graphidaceae and unrelated to Fissurina and
instead close to Coi !i,;.i. _, .i *;, *. ii
Acanthothecis poitaeoides (Nyl. ex Tuck.) E. A.
Tripp & Lendemer-Figure 4C (Seavey &
', .,1ey 10843). Species of .. .ii. ii' ,,h ,i are
recognized by their non-carbonized lirellae,
warty paraphysis tips and usually non-amyloid
ascospores. A taxonomic treatment of North
American species is given by Tripp et al.
(2010).
Actinoplaca spec.-Figure 4D (Sanders 10504.1,
1 014-/.6). This material ,.t .- *-.tl.. an
undescribed foliicolous per-i- most probably
belonging in Actinoplaca (Lul.Mv;t 2008).
It produces abundant, very shortly stalked
hyphophores that carry a rounded mass of
agglutinated diahyphae interspersed with algal
cells. No apothecia were found in the material.
*Amandinea endachroa (Malme) Marbach-
Figure 4E (Breuss _"'.'36*). Species of
Amandinea are treated in Marbach (2000). The
genus is probably heterogeneous and needs a
revision using molecular methods.
Amandinea punctata (Hoffm.) Coppins &
Scheid.-Figure 4F (Seavey & Seavey
10712*).
Anisoineridium ambiguum (Zahlbr.) R. C.
risris-(Harris & Buck s.n.).
Anisoineridium aureopunctatum R. C. Harris-
(Buck 5-i-4."). Harris (1995) provided a key to
Anisomeridium species, both for Florida and
world-wide.
Anisoineridium phaeospermum R. C. Harris-
(Lendemer 15538).
*Anisomeridium subnexum (Nyl.) R. C. Harris-
Figure 4G (Liicking & Rivas Plata 26616).


Anisomeridium subprostans (Nyl.) R. C. Harris-
igure 4H (B. ,.!ii,;: 7706, Seavey & Seavey
10700).
Anisomeridium terminatum (Nyl.) R. C. Harris-
(Lendemer 15516).
Anisomeridium spec.-Figure 5A (iT,. ,'l: &
Rivas Plata 26598*, 2661 6! The material
agrees with Harris 29392A in Harris (1995) and
probably represents an undescribed species.
Unfortunately the material is not abundant
enough for a formal description.
ml,,,. oith. .Ma nanum (Zahlbr.) R. C.
Harris-(Harris & Buck s.n.). Species of
Anthracothecium are treated in Harris (1995)
ialr,,,tn ifJ' ; Uprasinum (Eschw.) R. C.
Harris-Figure 5B-D (Beeching 7583, Lay
09-0036, L.._l,_r & Rivas Plata 26591*,
26595*, Mfercado-Diaz 389, Nelsen 4095,
Seavey & Seavey 10602).
*Arthonia antillarum (Fee) Nyl.-Figure 5E-F
(Common 7327E, Mercado-Diaz z //') The
genus Arthonia is not well-understood and
the present study revealed several additions
to the North American lichen checklist;
some of these are widespread in the coastal
plain but have not yet been reported in the
literature. The taxonomy of the pr-; found
here is summarized in Table 2. Compare also
Coniarthonia.
Arthonia cinnabarina (DC.) Wallr.-Figure
5G-H (Common 7-ij^1C, Crane ILLS 60432,
Seavey & Seavey 10611*). As pointed out by
Grube (2007), this is a difficult species complex
with variation in ascomata morphology and
ascospore size and septation. In our material,
Common 7i'lC has few marginal pruina and
the ascospores are mostly 4-septate and 20-25
x 6-8 pm.
iirdi,ria complanata F6e-Figure 6A (Lay 09-
004i1, Seavey & Seavey 10542).
Arthonia ilicina Taylor-Figure 6B (Mercado-
Diaz 410). The material fits well with the
description given in Grube (2007), particularly
the apothecia being first immersed and covered
with remnants of bark tissue resembling a







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,' 1,


Table 2. Simplified keytable to identify species ofArthonia cited in this work (only lichenized species included). Septa = number
of ascospore septa: muri = muriform; Shape = ascospore shape: iso = isolocular (cells equal or middle cell sometimes slightly
enlarged), macro = ii ,c L-..-c pii c.- (upper and sometimes also lower end cell .- oi ,,,- ,it submacro = submacrocephalic (second
upper cell enlarged); Ascomata = .11. > ,, of ascomata: irreg = irregular; Color = color of ascomata; Pruina = I ... i i- cover of
ascomata; Emergence = emergence of ascomata; Chemistry = secondary thallus chemistry (not including ascomata pigments):
lichexan = ]h.. -..i lii.i. psor = psoromic acid; Other = other characters: inspersed = inspersed hymenium, K+ pigments =
crystallized pigments present in excipulum, epithecium and/or hymenium, segmented = lirellae divided into segments; Size =
.. I ,, -, size in itm :I., I i d1 upper range for length and size; the typical lower range is about 30% less). Detailed hymenial
and ascus reactions with iodine solution are not given for lack of space (consulting specialized literature for Arthonia species
is highly recommended; e.g. Grube 2007).


Species of
Arthonia

A. antillarum

A. aff. pruinata

A. redingeri

A. rubrocincta

A. cinnabarina

A. speciosa

A. wilmsiana


A. rubella

A. pruinosella

A. simplicascens

A. complanata

A. ilicina

A. platygraphidea

A. cyrtodes

A. interveniens

A. macrotheca

A. mesoleuca

A. mirabilis


..p.i Shape Ascomata Color Pruina .. .. ....'- hi,... Other Size


4

3

3-4

4-7

3

5-7

5

5-7

11-17

11-13

muri

muri

muri

muri


submacro

submacro

macro

macro

macro

macro

submacro

macro

macro

submacro

submacro

macro

macro

+ iso

+ iso

+ iso

iso

iso

iso


irreg-lirellate

rounded-irreg

n o-- I I I.
i11 li '11n.! .


irreg

lirellate

irreg

lirellate

lirellate--11.I i. -

irreg

rounded-irrI.

rounded-irreg

rounded-irreg

rounded-irreg

rounded-irreg

rounded-irreg

rounded-irreg

rounded-irreg

n ,. -~.'I I l '


pale adnate hiI..- .. ,i

pale pruinose erumpent -

cinnabar pruinose erumpent -

cinnabar pruinose erumpent psor

cinnabar pruinose erumpent -

brown-orange erumpent -

dark red adnate -

brown adnate -

brown adnate -

black pruinose adnate -

black adnate -

black adnate -

brown-black bark erumpent -
tissue
black adnate -

black adnate -

black adnate -

black adnate -

black adnate -

brown-black adnate -


18 x 5

28 x 10

K+ pigment 18 x 6

K+ pigment 22 x 6

K+ pigment 28 x 8

K+ 4.jIc41 20 x 8

K+ pigment 25 x 12

segmented 18 x 6

30 x 12

15 x 5

15 x 6

5 x 8

35 x 12

70 x 20

inspersed 60 x 15

30 x 10

inspersed 70 x 30

K+ ';.2-ii c.1 50 x 20

K+ pigment 55 x 20






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


coarse pruina and later becoming erumpent to
almost sessile and ,I;;, ,;I,. I/,, I;ii-
**Arthonia interveniens Nyl.-Figure 6C-D
(Common 7303A, 7.:' ,A).
Arthonia macrotheca F6e-Figure 6E-F
(Beeching 7597, Lendemer 15515, Seavey &
Seavey 10167*).
Arthonia mesoleuca Nyl.-(Lendemer 15544). The
description for the North American material
of this species in Lendemer et al. (2009b) is
reminiscent of another, recently described
pfei-; A. mirabilis Grube, which was also
identified from Fakahatchee material. The
main difference is in ascospore size (less than
50 j[m long in A. mesoleuca versus more than
50 pm long in A. ,ii 1,;i;-) and the often more
lirellate ascomata in A. mirabilis; however, it
is possible that both are conspecific, in which
case A. mesoleuca would be the older name.
Arthonia mirabilis Grube-Figure 6G-H
(Common 7265).
"AidJilwii" montagnei (Tuck.) R. C. Harris-
(Buck 54466). This name is here applied
to a species that apparently belongs in
the genus Ci.J ',". i.. i' (and therefore not
included in Table 2), with 8-spored asci
and 2-O-methylperlatolic and confluentic
acids. This taxon has been confused with the
recently described Cryptothecia i. ; ',Li..i. w.i-.
(Seavey 2009), which also has 8-spored
asci but smaller ascospores and contains
psoromic acid and lichexanthone. The name
( n ../'..._., i t,, /.. .. Tuck. (Tuckerman
1872) is apparently based on material that
Montagne (1842) first identified as atypical
( ./~. /. t, -i lacteum Fee, based on an Auber
collection from Cuba. We have not yet seen
that material and in the light of the chemical
,i <,.- i j, of C:)1 di.. .i.. species, it is possible
that the Auber material is not conspecific with
the taxon from southeastern North America
named here A i -,ii ,i( it .,-:. i. in which
case a new species name would be required.
For this reason, we have refrained at this time
from simply transferring Arthonia m. 'ii,:..:;..


to 0 ) , ;', t,
Arthonia aff. pinastri Anzi-Figure 7A (Lay
W~/ 1 /2A). We have not yet found a definite
name for this taxon which has macrocephalic,
3-septate ascospores about 15 x 5 pm in size
and abundant pycnidia and appears to be related
to the bulk of foliicolous taxa (LOcking 2008).
The material fits A. pinastri in most details,
including the abundant pycnidia (Grube 2007),
except that it is lichenized.
Arthonia plite ; aphiq/J,.i Nyl.-Figure 7B
(Jnl..Jl.- & Rivas I'iLi..1 26583*, 2?:4"-*,
Seavey & '., ,'ey l,.It '. i).
Arthonia aff. pruinata (Pers.) Steud. ex A. L.
Sm.-Figure 7C-D (Common 7372A). This
material deviates from typical A. pruinata
(Grube 2007) by its larger ascospores (24-30
Ipm).
Arthonia pi in.1,i-l i Nyl.-Figure 7E-F
(Common 7327A).
Arthonia pyrrhuliza (F6e) Nyl.-Figure 7G-H
(Lay 09-0017A, Mercado-Diaz 438b).
Arthonia redingeri Grube-Figure 8A-B
(Common 7259U).
Arthonia rubella (Fee) Nyl.-Figure 8C (Licking
& Rivas Plata 26586*).
Arthonia rubrocincta G. Merr. ex Grube &
Lendemer-Figure 8D (Buck 54444, Lay 09-
I', i2B, Lendemer 15611, Licking & Rivas
Plata 26582*, Nelsen 4010, Seavey & Seavey
11001). This name was recently validated
(Grube & Lendemer 2009).
**Arthonia simplicascens Nyl.-Figure 8E-F
(Common 7291D). We apply this name as it
fits the type specimen from Sri Lanka rather
..iI in ascomata morphology and anatomy
and ascospore type. However, it is likely that
there are additional names available for this
relatively non-descript taxon.
Arthonia speciosa (Miill. Arg.) Grube-Figure
8G-H (Common 72911, Liicking & Rivas Plata
26622).
Arthonia spec.-(Lay 09-0191). This is an un-
named, yet characteristic taxon. The ascomata
are black, rather plane, and irregular in shape;
the hamathecium is slightly inspersed with oil






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


droplets and reacts K- (no crystalline pigments
present); ascospores are muriform, about 35-
40 x 10-15 pm in size and become brown when
mature. Arthonia macrothecia is most similar
but has much larger, hyaline ascospores. The
African A. 7-.. v.,..ir.. Nyl. also resembles this
material but its ascospores are persistently
hyaline and have an enlarged distal cell.
Arthopyrenia cinchonae (Ach.) Miill. Arg.-
Figure 9A (Lendemer 15615A). Species of
Ai i,, ') i, ii. t are keyed out in Harris (1975,
1995).
Arthopyrenia lyrata R. C. Harris-Figure 9B
(Harris & Buck s.n.).
Arthopyrenia planorbis (Ach.) Miill. Arg.-
Figure 9C (Seavey & Seavey 10165).
Arthopyrenia aff. planorbis (Ach.) Mill. Arg.-
Figure 9D (I..h lw; & Rivas J'/,..; 26787b).
This material agrees with A. planorbis in
ascospore type but has consistently excentric
(lateral) ostioles. We have not found a name for
this taxon but refrain from describing it as new,
since the genus is in further need of revision
and possible epithets might be hidden among
other generic names.
Arthothelium spectabile A. Massal.-Figure 9E
(Breuss -'. -0*, Liicking & RivasPlata 26576*,
26581). This species strongly resembles an
Arthonia with muriform ascospores, but differs
in the strongly compacted, stromatic structure
of the hymenium in which the asci are formed
in locules and surrounded by a delicate network
of anastomosing paraphysoids. The entire
hymenium in the present material appears
inspersed.
Arthothelium spec.-(7/./,ji_;u; & Rivas I'/;..i
26500). The scanty material does not fit any of
the described I....- ; -, in the genus.
*Aspidothelium cinerascens Vain.-Figure 9F
(Harris & Buck s.n.). For key to species of
A .pi. ,,1,, 1h i:,,ii see Lucking (2008).
S*A'< iJhcI.'li.un geminiparum (Malme) R.
Sant.-Figure 9G (Lendemer 26576, Licking
& Rivas Plata 26600c).
*Aspidothelium scutellicarpum Liicking-Figure
9H (Breuss 28787*).


Astrothelium confusum Miill. Arg.-Figure 10A
(7n~._/, ,;- & Rivas I'i;a.. 26691, Seavey &
Seavey 10187). This genus is treated in Harris
(1995). Molecular data suggest that the genus
concept is artificial and most probably Harris'
(1995) idea of including the bulk of species of
A h ,iin iil,,ii, C. m & i, , i, C; i /l-,,, .
Laurera, and !ip ,.. e/i.. I/, within a single
genus Laurera (or Bathelium) is correct (Nelsen
et al. 2009). In addition, the t ... i;, concept
in the ziA .., ,ii, ia,: variolosum /2IT /. ;h / ,,ih
nilidiusculum a' r,'n.it,: needs revision. We
do not agree with Harris (1995) that species
with either fused or separate ostioles should be
lumped, as there are substantial differences in
other morphological features as well. In addition,
hymenial ;irf,;,, has not yet been explored
as a taxonomic character in this complex,
even if its use is accepted for other genera.
*Astrotheliuim JIiI'' ,i!,lk Miill. Arg.-Figure
10B (Licking & Rivas Plata 2662 7*, 26630*).
This tp ... i<, is quite distinctive due to its very
large ascospores (70-85 x 17-22 ipm) which,
however, have only 5-7 septa with distinctly
diamond-shaped lumina, thus looking like
gigantic ascospores of the Ti i Ihi,. iii-type.
The thallus and especially the perithecial
warts, which contain 3-6 aggregate perithecia
with fused ostioles, are UV+ golden-yellow
(lichexanthone). The species was described
from Cuba (Muller Argoviensis 1885), so its
presence in Florida is not a surprise.
Astrothelium diplocarpum Nyl.-Figure 10C
(... h./~'- 7626, Buck 54444, Lay 09-00i:/,
Nelsen 4012, Seavey & Seavey 10087).
Astrothelium galbineum Kremp.-Figure 10D
(Crane ILLS ': i 55, Licking & Rivas Plata
26596, Nelsen 4013).
Astrothelium variolosum Miill. Arg.-Figure
10E-F (Beeching 7,-io, Breuss _.".'.'*, Lay
09-01 ', Licking & Rivas Plata 26691*,
26771*, Nelsen 40lJ, Seavey & Seavey
10092). This taxon is supposed to have a clear
hymenium, but Harris (1995) also included
specimens with inspersed hymenia. We found
one collection with an inspersed hymenium






LUCKING ET AL.: Lichens of Fakahatchee t.i i Preserve State Park


(Breuss ''- /) and suspect that this represents
a separate taxon, but no name appears to be
available for this form.
Aulaxina microphana (Vain.) R. Sant.-Figure
10G (Sanders 10521.1). Licking (2008)
provided a key to -p I....-. i-- of Aulaxina.
Aulaxina ,jiiaildsii: li (Stirt.) R. Sant.-Figure
10H (Sanders 10521.7).
Bacidia :.,'; i.ih. ai, Malme.-Figure 11A
(Seavey & Seavey 10095*). Species of Bacidia
can be identified using Ekman (2006).
Bacidia campalea (Tuck.) S. Ekman & Kalb-
Figure 11B (Breuss 28735*, .- '-"' Seavey &
Seavey 10i045).
Bacidia heterochoa (Miill. Arg.) Zahlbr.-Figure
11C (Lay 09-o-),,, Ltlr. & R `,,. Ti';,.
26621*, Seavey & Seavey 10234).
Bacidia aff. heterochroa (Miill. Arg.) Zahlbr.-
Figure 11D (Licking & Rivas Plata 26610).
This material differs from Bacidia heterochroa
in the gray-black apothecia with flat disc and
gray margin.
Bacidia hI otdh,:c ,idli (Nyl.) Zahlbr.-Figure 11E
(I A in- & Rivas Plata 26605, 26606, 26607*,
26608, 26609).
Bacidia aff. ho,,Ith, l ',l (Nyl.) Zahlbr.-Figure
11F (Licking & Rivas / !'., 26600b). This
material keys to B. I'.1....i. l in Ekman
(1996) but differs in the dark apothecia without
orange tinge and the persistent margin.
Bacidia medialis (Tuck.) Zahlbr.-Figure 11G
(Buck 54433, Ln ..lI,? & TR w,. Plata 26601*,
26603*, Mercado-Diaz 433).
Bacidia aff. medialis (Tuck.) Zahlbr.-Figure 11H
(Licking & Rivas Plata 26606). This material
resembles Bacidia medialis but has darker
apothecia and the internal parts (excipulum)
have an orange, K+ purple band. The material
is also similar to B. h, ilh, ;, -, /. in which the
excipulum is pale orange and not K+ purple.
Bacidia mutabilis Malme.-Figure 12A (Licking
& Rivas Pil.u, 26603, 2,1:i4*, Seavey &
',. .. ey 10129).
Bacidia russeola (Kremp.) Zahlbr.-Figure 12B
(Breuss f2*. ", ',, Lay 09-0019, Mercado-Diaz


Bacidia schweinitzii (Fr. ex E. Michener) A.
Schneid.-Figure 12C (Seavey 13331*).
Bacidia aff. schweinitzii (Fr. ex E. Michener)
A. Schneid.-Figure 12D (Liicking &
Rivas Plata 26602a). This material covers
the taxon with brown apothecia and brown
epithecium included by Ekman (1996) in
Bacidia schweinitzii. In our view, this is a
taxon separate from B. schweinitzii s.str. with
black apothecia and red-black hypothecium,
but there appears to be no alternative name
available. The type material of all synonyms
cited under B. schweinitzii by Ekman (1996)
seem to represent the typical form with black
apothecia. We have refrained from formally
describing the taxon with brown apothecia
since our material is moderatly developed and
segregating this taxon requires restudy of the
many specimens cited by Ekman (1996).
i:tridin spec. 1-Figure 12E (Licking & R'i.,
PF,,,i/ 26611). This material does not key to
any species in Ekman (1996). The apothecia
are large, brown and irregular in outline, with
an orange excipulum and hypothecium. The
ascospores are 40-50 x 4-5 [im and thus
unusually broad, with 5-11 septa. Most similar
in ascospore type are B. insularis Zahlbr. and B.
subincompta (Nyl.) Arnold, which have black
apothecia with different I"' tit, i i I pigments
internally (Ekman 1996).
Bacidia spec. 2-(Lay 09-0213). The taxonomic
status of this material is unclear. The granular,
green thallus and the more or less iii.ir.it., t
turbinate to convex, dark gray-brown apothecia,
together with the 3-septate ascospores, ...; -.t
a species ofBiatora orMycobilimbia, but none
of these two genera includes anything similar.
Biatora tetramera (De Not.) Coppins agrees in
general morphology but has larger ascospores.
Since the ascospores of the present material
are rather narrow (to 13 x 2.5 rpm) and the
excipulum consists of parallel hyphae, Bacidia
is a likely home, but no matching species was
found in the key provided by Ekman (1996).
Bacidina varia S. Ekman-Figure 12F (Lay 09-
ioo; I).






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1h


Bactrospora denticulata (Vain.) Egea &
Torrente-Figure 12G (Licking & Rivas Plata
26737b, Seavey & Seavey 10325). The two
species of Bactrospora found at Fakahatchee
are characterized by cylindrical ascospores
lacking constrictions and a K/I-, rather
thick excipulum (Egea and Torrente 1993).
Bactrospora denticulata has a denticulate
apothecial margin and ascospores to 130
jim long, whereas B. myriadea has a smooth
margin and ascospores less than 100 um long.
r-:ri,.'n,. ti,, rn! ,.ia. :. (F6e) Egea & Torrente-
Figure 12H (:.'" :- 5/ i' Lay 09-0 / i, 09-
0212, Seavey & Seavey 10326).
Wli, va iJifr, curtisii (Tuck.) Marbach-Figure 13A
(Seavey & Seavey 10432). Marbach (2000)
provided a key to P. ,I ,'if,. i species.
ir,,di rjt! imshaugiana (R. C. Harris)
Marbach-Figure 13B I,' /, ,.. 28890*).
Bathelium :,naliieriininr (Tuck.) R. C. Harris-
Figure 13C (Liicking & Rivas Plata 26617,
Nelsen 4149). Species of Bathelium are treated
in Harris (1995).
Bathelium ,nr 'l,{nrri,; ri,,' (Eschw.) Trevis.-
Figure 13D (Lay 09-,0/ 17).
Ir,'i,,;in,, leucoxantha (Spreng.) R. Sant.
& Hafellner.-Figure 13E (;':,.. :'1:"*,
Hodkinson 10553, Nelsen I'~-).
Buellia stillingiana J. Steiner-(?',, hir in 7615).
*Byssoloma chlorinum (Vain.) Zahlbr.-Figure
13F (,.,/.. , 4078, %.,;p/,,' ,- 135). Licking
(2008) provided a key to neotropical Byssoloma
species including those found in Florida.
Byssoloma leucoblepharum (Nyl.) Vain.-Figure


13G (Li/cking & Rivas Plata 26692*, Seavey
& Seavey 102 i'.).
Byssoloma meadii (Tuck.) S. Ekman-Figure
13H (Buck 54387, Lay 09-0172, Lendemer
15572, Licking & Rivas Plata 26693, Seavey
& Seavey 1005 i).
Byssoloma subdiscordans (Nyl.) P. James-
(Safi'anek 131).
Calicium 1 p,'ri-Lhi, ,E' Nyl.-Figure 14A-B
(Harris & Buck s.n.).
*e ia IqE rihi ,-,a /ioVezda ex Chaves & Liicking-
Figure 14C (Common 7322B). This recently
described species (Lumbsch et al. 2011) is
closely related to C. ,, ....., but differs in the
heavily pruinose apothecia. In contrast to
C.l.',i.,i,. < floridana, the apothecia disc is
orange-brown and the pruina yellowish, and
also the apothecial margins are pruinose.
***Calopadia fl.uidiLi,l Hodges & Liicking-
Figure 14D-G (Hodges s.n.). See p. 138 for
description of this new species. The species of
C.,, ..' .i,. currently known from Florida are
summarized in Table 3.
Calopadia fii, (; (Miill Arg.) Vezda-Figure
14H (Lay 09-0186, k., p ,,A-i 72, 73, Seavey &
Seavey 10365*).
***Calopadia imshaugii Common & Liicking-
Figure 15A-F (Common 7322F). See p. 139
for description of this new species.
*Calopadia I.-,utur,.Ilr a (Nyl.) Kalb & Vezda-
Figure 15G-H (Lay 09-0219, Liicking & Rivas
Plata 26696, Nelsen i11 71).
*i al2rIqi p,'l'iTf/ilei (Nyl.) Vezda-Figure 16A
(Nelsen ii a).


Table 3. '- nlphi. I- keytable to identify species of Calopadia cited in this work. Ascospore length in tim.

cl... ;- of Disc color Pruina H. .h.ii1 ii'M color .'.. ..'.. number Ascospore length
Calopadia
C. imshaugii orange-brown white-gray brown (1-)2(-3)/ascus 50-80
C. ...., '.' orange-brown yellowish light brown single 80-110
C. editae orange-brown yellowish light brown single 50-80
C.floridana gray-brown white-gray dark brown single 70-80
C. fusca orange-brown light brown single 60-85
C. puiggarii gray-brown dark brown single 55-85
C. lecanorella black dark brown single 60-80
C. subcoerulescens black i.i-.., single 55-85






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


Calopadia puiggarii (Miill. Arg.) \ 7.A.-FI'igure
16B (Beeching 7659, Crane ILLS 60444, Lay
09-0005, Lendemer 15616, Licking & Rivas
Plata 26695*, 26695, -., i ,.. 76, 81, Seavey
& Seavey 10367).
*Ct',l ~ien subcoerulescens (Zahlbr.) Vezda-
Figure 16C (Buck 54. :, ').
Cadhqidia spec.-(Common 7399D). This
material bears only campylidia and cannot be
identified to species level but is distinctive due
to its conspicuously verrucose thallus, a feature
uncommon in C.i,.y..;..i. and the related
genus Ti-,,,, i.. The campilidioconidia
associate with photobiont cells, a feature more
commonly observed in the genus ;, !,p,'ji,,i .
(Lucking 2008).
(' alrps1a, a -inrihra (Tuck.) Zahlbr.-Figure 16D
(Lucking & Rivas Plata 26568b). This is one of
the brown Caloplaca species, characterized by
the pruinose apothecia resembling those of a
Gyalideopsis (Wetmore 1994).
(_alpla.:'n a epiphora (Taylor) C. W. Dodge-
Figure 16E (Beeching 7688, Buck 54398,
Common 7310, Lay 09-0052, Lendemer
15576, Seavey & Seavey 1/ ; 13*). This species
is characterized by the orange and densely
isidate thallus.
(GalDll.' a hoI,:, .rajn (Hoffm.) A. E Wade-
Figure 16F ( i;, ,-iu." & Rivas Plata 26698).
This material features mostly deformed
apothecia with brownish color, but some
apothecia with orange color and flat discs are
present corresponding to those typically found
in this species.
( arinu.r'hal cryptochlorophaea (Hale) Elix &
Hale-(. _.. ',~h;i:,. 7612, Mercado-Diaz i-',
Seavey & Seavey 10056).
(Ci_, 'rn.. ',fa salicinifera (Hale) Elix & Hale-
(Harris & /.','-/ s.n.).
Catillochroma endochroma (F6e) Kalb-Figure
16G (Beeching s.n.*). This material was
first believed to represent a new species in
'i /,/, .',hi. I/, since the ascospores appeared to
lack septa. However, some 1-septate ascospores
were eventually found, and the apothecial
anatomy, with the outer excipulum consisting of
radiating cell rows, the inner excipulum being


of medullary structure, and the hypothecium
being pale, as well as the aeruginous epithecium,
characterize the material as Catillochroma
(Kalb 2007). Species of .1 ,i,'n,.. iL can be
remarkably similar in appearance, especially
as Malcolmiella granifera and related taxa also
have a medullary excipulum, but apart from the
non-septate ascospores, these *.- ... ;- differ in
the outer paraplectenchymatous excipulum and
the brown-black hypothecium. Catillochroma
and Lopezaria have been synonymized with
Megalaria (Fryday & Lendemer 2010). We do
agree that the limits between the three genera
are not clearcut but differences between the
type species, including in ascus structure, exist.
Unfortunately, no sequence data are available
for Catillochroma; analysis of the available
sequences of the large subunit of the nuclear
ribosomal DNA (nuLSU) of one specimen
of 3L *:..din,( gross and two specimens of
Lopezaria versicolor (Lucking, unpubl. data)
show that both are in the same clade within
Ramalinaceae, but on very long branches
each, which supports a close relationship but
not congenerity, considering that nuLSU is a
relatively conserved gene partition and hence
species within the same genus are expected
to appear on short branches. Therefore, we
maintain the genera separate pending further
studies. The overlap of anatomical and
chemical features between species assigned to
the different genera documented by Fryday &
Lendemer (2010) is not necessarily evidence to
merge them, since phylogenetic studies show
that genetically distinctive clades can evolve
similar phenotypic characters in parallel (1'- .]i.
Plata et al. 2011).
Celothelium aciculiferum (Nyl.) Vain.-Figure
16H (Common 7396D, Lay 09-01'" i)
(Cj'd.c'(endica brunneola (Ach.) Miill. Arg.-
Figure 17A (Beeching 7613, Buck 54451, Lay
09-1-'4', Lendemer 15630, Seavey & Seavey
10058*).
C hhmciL, ti a floridaia R. C. Harris-(Beeching
7663, Crane ILLS 60436, Seavey & Seavey
10435*).
*(. hlapp, chionostoma (Nyl.) Rivas Plata &






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


Mangold-Figure 17B (Common 7321A). The
material agrees perfectly with the type of this
species in the morphology of the thallus and
apothecia, which are somewhat intermediate
between Ch,7 q,* / and Thelotrema, as well as
the brown excipulum and hardly discernable
periphysoids. The ascospores are, however,
larger than in the type and material from Costa
Rica (25-30 ipm versus 15-20 !tm) and with
more numerous septa (9-13 versus 5-9).
Since only few specimens are known, we
refrain from considering the Florida material a
separate taxon, until the variational amplitude
of the Florida populations has been established
in I, ir'i',i -.tI with the material from Costa
Rica and Panama (Hale 1978).
Chapsa Iplati -apa (Tuck.) A. Frisch-Figure
17C (Beeching 7691, Lay 09-0053, Lendemer
15646, Licking & Rivas Plata 26573*,
26574*, Seavey & Seavey 1~."-/*). This
species is commonly found on the thin trunks
and branches of swamp dogwood.
*Cihapi platycarpoides (Tuck.) Breuss &
Liicking-Figure 17D (Breuss 2''*,
Lacking & Rivas Plata 26571*). See p. 140 for
new combination and discussion.
** (iCh, ha .M tIbpa ,'itn (Hale) Mangold-(Common
7 i",.i). The material fits the type from Sri
Lanka well in the endoperidermal, pale thallus,
the irregular apothecia, the amyloid ascospores
with rather thin septa, and the lack of secondary
substances. Previously known from Sri Lanka
and Australia (Hale 1981; Mangold et al.
2009); new for the New World.
Chrysothrix xanthina (Vain.) Kalb-Figure 17E
(Beeching 7591, Crane ILLS 60463, Buck
54437, Lay 09-0055, Lendemer 15592, Nelsen
lt, .u, Seavey & Seavey 10060).
(',inrni beaumontii (Tuck.) Fink-(Harris &
Buck s.n.).
('I,hmni, ,a-Ji-,pr.,-ia (Flirke ex Sommerf.)
Spreng.-(1F.Z ,_:,/,, 7712, Lay 09-0056,
Lacking & R',, .. Plata 26699*, Mercado-Diaz
415, Seavey & Seavey 10193).
(',iJani didyma (Fee) Vain.-(Crane ILLS
*: 1i76, Lendemer 15673, Seavey & Seavey
10063).


Cladonia didyma var. vulcanica (Zoll. & Moritz)
Vain.-(Harris & Buck s.n.).
Cladonia ira ',-'hi Tuck.-(Lendemer 15665).
Cladonia subradiata (Vain.) Sandst.-(Z,.', :-;/
7710, i,, 15641, Crane ILLS 60453,
H, '.i, .,,1 10541, Lendemer 15600, Licking
& Rivas Plata 26700*, Mercado-Diaz 414,
Seavey & Seavey 10185*).
Clathroporina isidiifera R. C. Harris-Figure
17F (Beeching 7662, Licking & Ri,... Plata
26768, Nelsen 4124, Seavey & Seavey 10 :').
C/7,i :luv*,;;i,., comprises species of Porina
s.lat. with thallus-dominated perithecial
verrucae and shiny thallus and (dark) prothallus;
the Florida species are treated in Harris (1995).
Cift,~nnl,,,ri, subpungens (Malme) R. C.
Harris-Figure 17G (Lay 09-0129, 09- ".'
Lacking & Rivas Plata 26758, Seavey &
Seavey 10241*).
Citilftnnlo;,,a tetracerae (Ach.) R. C. Harris-
Figure 17H (Licking & Rivas Plata 26767,
Seavey & Seavey 10242).
Coccocarpia Jim.d/:n.'lli \ Vain.-Figure 18A
(Harris & Buck s.n.).
Coccocarpia ery'r/ifi 'ht (Spreng.) Swinscow &
Krog-Figure 18B (Lendemer 15508, Nelsen
f,,,,, Seavey & Seavey I '.)
Coccocarpia palmicola (Spreng.) Arv. & D. J.
Galloway-Figure 18C-D (Crane ILLS 60441,
Lay 09-0057, Lendemer 15632, L#cking &
Rivas Plata 26701, Seavey & Seavey 10383*).
*Coenogonium congense C. W. Dodge-Figure
18E (Lay 09-013 i). The genus Coenogonium
(including Dimerella) is treated with a world-
wide key including all Florida species by Rivas
Plata et al. (2006).
*Coenogonium geralense (P. Henn) Liicking-
Figure 18F (Lay 09-0135, iilh. i,4' & RIo,.
FP. i,/ 26702).
Coenogonium interplexum Nyl.-Figure 18G
(Lay 09-0132).
Coenogonium linkii Ehrenb.-Figure 18H (Lay
09-0002, Nelsen i t i )
*Coenogonium luteocitrinum Rivas Plata,
Liicking & Umaiia.-Figure 19A (Lay 09-
0131).
Coenogonium luteum (Dicks.) Kalb & Liicking-






LUCKING ET AL.: Lichens of Fakahatchee i.i i Preserve State Park


Figure 19B (.-, ;' hi1u 7614, Lendemer 15589,
Seavey & Seavey 10130).
*Coenogonium subdentatum (V6zda & G. Thor)
Rivas Plata, Liicking, Umana & Chaves.-
Figure 19C (Lay 09-01.,-
*Coenogonium mli,!u, : i,',eu (Vezda &
Farkas) Liicking, Aptroot & Sipman.-
Figure 19D (Licking & Rivas Plata 26703*).
('Ith-m., fiirfira,-,'cun (Arnold) Du Rietz-
(Harris & Buck s.n.).
( i A/,- i ptli'l luiw Ach.- (-.:,',' I'n, 7713,
I, /. 1 1,, 11 3015).
*C(4 mi',rtinia wilmniana (Miill. Arg.) Grube-
Figure 19E (Beeching 7702, Common 7291G).
Cratiria lauricassiae (Fee) Marbach-Figure 19F
(Common 7_ '-E, 7329C).
Crocynia gossypina (Sw.) A. Massal.-Figure
19G (Seavey & Seavey 10473*).
Crocynia t\Eviil,'~ Nyl.-Figure 19H (Beeching
7592, Crane ILLS 60454, Lay 09-i' ~,-
L#cking & Rivas Plata 26705*, Nelsen 4045,
Seavey & Seavey lo,;, i).
*Crvpid,-ii nana (Tuck.) D. Hawksw. &
Dibben-Figure 20A (Breuss 28711).
*Cryptothecia effusa (11II. Arg.) R. Sant.-
Figure 20B (L#cking & Rivas Plata 26683).
The otherwise well-developed material has
mostly sterile ascigerous parts but in two
..pi' ii, n .. a few young asci where found with
eight undeveloped ascospores per ascus.
Cryptothecia evergladensis Seavey-Figure
20C-D (Lay 09-0012, L#cking & Rivas Plata
26670).
***Cryptothecia miniata Vain. ex Liicking-
Figure 20E-F (Lay 09-0063, Licking &
Rivas Plata 26678, 26679*). See p. 140 for
description of this new species.
*Cryptothecia ptIe..:i,,. 'di,,tu Sparrius.-
Figure 20G (Beeching 7641, L#cking & Rivas
Plata 26680, Nelsen /i -'?). In this material and
other -..pt I i r,. n.. from the Neotropics, the entire
thallus contains gyrophoric acid and reacts C+
red, not just the soralia as in the paleotropical
populations (Sparrius & Saipunkaew 2005).
Cryptothecia striata G. Thor-Figure 20H
(Beeching 7678, Crane ILLS 60439, Lay 09-
0009, Lendemer 15;'iO, Licking & Rivas


/i', 26685*, 26687, Mercado-Diaz .','
Seavey & Seavey 10066). This species is often
misinterpreted in the literature as having isidia.
Isidiate with a chemistry similar to that of C.
striata (gyrophoric acid) must be referred to
H,. I'. ;l',/iti. philippinum (Vain.) Aptroot &
Lacking (Aptroot et al. 2009). Cryptothecia
striata is one of the most common and
widespread species of the genus in the
Neotropics and consistently fertile, always
lacking isidia.
*Dictyonema phyllogenum (Miill. Arg.)
Zahlbr.-Figure 21A (Common 7363D).
*Dictyonema sericeum f. phyllophilum Parm.-
Figure 21B (Harris & Buck s.n.).
Diorygma junghuhnii (Mont. & Bosch) Kalb,
Staiger & Elix-Figure 21C (Common 7418D,
7425A, Lendemer 15639). This genus has been
monographed by Kalb et al. (2004) and North
American t 1..-. ,, are treated in Tripp et al.
(2010).
***Diorygma microsporum M. CAceres &
Liicking-Figure 21D (Lay 09 ut', i, Lucking
& RI ,. Plata 2.',i-4). This species, described
in a separate paper (Lumbsch et al. 2011), was
previously misidentified as An in. 'i./, 'i,
., .:.-,:,, (Nyl.) Staiger (Caceres 2007),
with which it agrees in the small ascospores
and norstictic acid but differs in the absence
of hymenial inspersion and the indistinctly
corticate thallus. Sequencing data showed that
the species belongs in Di,; ..,. instead, with
which it agrees in the thallus and ascomata
morphology (Lumbsch et al. 2011).
Diorygma poitaei (Fee) Kalb, Staiger & Elix-
Figure 21E (Crane ILLS 60430, Seavey &
Seavey 10548*).
Dirinaria dc,'niJliti, (Afz.) B. J. Moore-Figure
21F (Beeching 7593, Lendemer 15606, Seavey
& Seavey 10621*).
Dirinaria confusa D. D. Awasthi-Figure 21G
( ..... lai' 7619).
Dirinaria leopoldii (Stein) D. D. Awasthi-Figure
21H (Seavey & Seavey 10632*).
Dirinaria papillulifera (Nyl.) D. D. Awasthi-
Figure 22A-B (..... ci/,, 7617).
Dirinaria picta (Sw.) Schaer. ex Clem.-Figure






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


22C (Crane ILLS 60462, Lendemer 15561,
Nelsen /' ?9, Seavey & Seavey 10633).
Dirinaria j, r,,fr !.:,_u'\ (Vain.) B. Moore-
Figure 22D (Seavey & Seavey 10636*).
DyT,'i.TJln ,i./ :.li (Ach.)A. Massal.-Figure 22E-
F / 1,'1:54397, Common 7356L, 7380E, Crane
ILLS 60427, Lay 09-0176, Lendemer 15521,
Licking & Rivas Plata 26505, Mercado-Diaz
443, Seavey & Seavey 10681).
Echinoplaca areolata Liicking & W. R. Buck.-
Figure 22G (Common 7'r..:B, Lay 09-0020,
Lendemer 15701).
Echinoplaca cf. ,-'l' p1 THi. Fee-(Sanders
I' -r i. 6b). The material is sterile but suggests
this species based on the verrucose thallus,
white setae and acute white hyphophores with
subapically inserted diahyphal bunches.
*F-.ifl, p.i/a leucotrichoides (Miill. Arg.) R.
Sant.-(Beeching 7660).
Echinoplaca aff. leucotrichoides (Miill. Arg.)
R. Sant.-Figure 22H (Crane s.n.). This
material represents an undescribed species,
but the single thallus is too small to describe
it formally. It agrees with the pantropical E.
leucotrichoides in the short hyphophores with
brown-black upper part, but contrary to the
latter, the diahyphal bunches include numerous
small algal cells with are dispersed with the
diahyphae. This appears to be the first species
i; F; 'in ; .. with this mode ofjoint dispersal
of mycobiont and photobiont (Lucking 2008).
Echinoplaca Im,,-.r-nif;era Kalb & \ 7,..ia-F'igure
23A (Sanders 1:i_ i lb).
Echinoplaca similis Kalb & VWzda-Figure 23B
(Breuss '.'."' ).
rlnir iarip ia anguinella (Nyl.) Redinger-
Figure 23C (Seavey & Seavey 10429*).
*I ig,',*'HIa leucocheila (i, i,) Licking, Serus.
& Kalb-Figure 23D (Buck 54463, Lendemer

***TF ,ru,, ,tW! v.v.zait!t, Common & Liicking-
Figure 23E (Common 7356A3). See p. 141
for description of this new species. Fissurina
sensu Staiger (", "') is a rather large genus,
with over 60 species (Lucking, unpubl. data).
Species concepts have so far been confusing


and the delimitation of species found in Florida
is summarized in Table 4.
***Fissurina analphabetica Common &
Liicking-Figure 23F-G (Common 73561,
7356N, 7377E, 7380L). See p. 142 for
description of this new species.
*Fissurina cingalina (Nyl.) Staiger-Figure 23H
(Common 7323F).
Fissurina columbina (Tuck.) Staiger-Figure
24A (Buck 54399, Common 73231, Lendemer
15512).
***Fissurina confusa Common & Liicking-
Figure 24B (Common 7356F, 7368G, 7380D).
See p. 142 for description of this new species.
**Fissurina crassilabra Mont. & Bosch-Figure
24C-D (Common 7356C, 73681, 7 :'. ,). This
S..... ..;, closely resembles Fissurina insidiosa
C. Knight and Mitt., but differs in the strongly
amyloid ascospores. Fissurina insidiosa has
been reported for Florida (Harris 1995), but the
specimens have to be rechecked for ascospore
amyloidity. Archer (2009) synonymized
Fissurina subcontexta under E insidiosa,
whereas Galloway (2007) kept them separate
but instead listed Fissurina eq. wq i ,i; i (Nyl.)
Nyl. and Fissurina ; j,,: .., C. Knight & Mitt.
as synonyms of E insidiosa. This suggests
some confusion about species concepts and
the corresponding type material. Fissurina
ti.;. .h i-. has labiate lirellae ('swollen lips'),
clearly seen in the picture of the type (Hayward
1977), whereas in E. ,::*1,; i- d .. and E. rugosa
they are fissurine with inconspicuous labia;
in addition, F .-, _.iili has a carbonized
.,_.ipulum and blackened lirellae. The
confusion about EI. ',i.. i Iit probably stems
from Staiger (21,,2), who used the example
of F comparilis to denote carbonized lirellae
in her scheme of classifying lirellae types
in Fissurina and also correctly depicted
the i..-i,_uluki, as apically carbonized in the
lectotype of F comparilis (Staiger 2002:133,
fig. 30), but in the description of the species
gave the excipulum as non-carbonized. The
type material of i:; ,.. on the other hand,
agrees well with Fissurina dumastii and







LUCKING ET AL.: Lichens of Fakahatchee -i.in i Preserve State Park


Table 4. Simplified keytable to identify species of Fissurina cited in this work and otherwise reported from Florida. ;..,. =
number of ascospore septa: muri = muriform; iodine = ascospore .lin0 1, dir.t chemistry = secondary chemistry; inspersion=
hymenium inspersion; excipulum = excipulum carbonization; lirellae = lirellae shape; other = other characters: verrucose =
thallus verrucose, endoperidermal = thallus ili1, I i, i.1.A i i size = ascospore size in im (typical upper range for length and
size; the typical lower range is about 30% less).


Species of Fissurina

E varieseptata

. inspersa

E subnitidula

F humilis

E tuckermaniana

F rufida

E insidiosa

F crassilabra

E aggregatula

F ... . .... . .





E analphabetica

F tachygrapha

F cypress

F columbina

F confusa

E subcomparimuralis

F nitidescens

F aff. elaiocarpa

E scolecitis

E mexicana

F cingalina

F egena

F incrustans


Septa Iodine Chemistry Inspersion Excipulum
7
7 -

3 inspersed

3 + carbonized

3 1+ carbonized

3 I+ carbonized

3

3

3 1+

3

3

3 (T+)

3

3 (1+)

3 (carbonized)

muri -

muri 1+ psoromic -

muri I+ psoromic

muri carbonized

muri (1+) -


mun I+- -


muri --

muri -

muri (1+) -

muri (+) -


Lirellae Other Size

fissurine cortex loose 15 x 5

fissurine-gaping cortex loose 15 x 6

roof-like 13 x 5

roof-like 20 x 8

roof-like 35 x 12

labiate 20 x 10

labiate-gaping vemicose 18 x 9

labiate-gaping vemicose 18 x 8

labiate-aggregate 15 x 6

pseudostromatic 18 x 6

fissurine-radiate 15 x 7

fissurine cortex loose 16 x 6

fissurine 15 x 7

fissurine-gaping endoperidermal 20x 9

fissurine-gaping ecorticate 100 x 25

roof-like 30 x 15

labiate .. I.ir... vermcose 30 x 18

erumpent-fissurine 22 x 8

chroodiscoid 18 x 8

labiate-gaping vemicose 25 x 15

labiate 28 x 12

pseudostromatic 35 x 12

fissurine-gaping 30 x 15

fissurine-gaping 20 x 10

labiate -.. -.i". 20 x 9






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P',I 1


should be considered a synonym of the latter,
thus adding F dumastii to the lichen biota of
New Zealand. The main differences between P.
insidiosa and E dumastii are the labiate versus
fissurine lirellae and the verrucose-bullate
versus smooth to uneven thallus.
Fissurina cypress (Miill. Arg.) Lendemer-
(Beeching 7643, Common 7268C, Seavey &
.'. .,ey 10456). This species is most certainly
not a genuine Fissurina, on the basis of its
large ascospores and general habit. It fits better
into the Diorygma-Thalloloma complex but is
set apart by its I- ascospores. Molecular data
are required to clarify the systematic position
of this taxon.
*Fissurina ,. *.
(Common 7276B, 7368Q). This -peri-; is close
to Fissurina incrustans, which differs in the
erumpent lirellae with distinct labia and .*- 0i; '-
disc, resembling Platythecium rather than
Fissurina (Staiger 2002). Fissurina. i. n., in
contrast, has immersed lirellae with roof-like
margins and concealed to slightly exposed
disc. However, intermediate forms do occur
and in fact this intermediate lirellae type is
quite common in several species of Fissurina,
such as Fissurina dumastii. North American
material with this type of lirellae has also been
identified as tissurina nitidescens (Nyl.) Nyl.
However, this is based on misinterpretation
of the type material (Staiger 2002). Graphis
nitidescens Nyl. is validly described in
Tuckerman (1888), who cited a collection from
Cuba (Wright 68) as the only collection (based
on Nylander 1886). Three collections with the
name are present in Nylander's herbarium:
Cuba, Wright 267b (H-NYL 7459!), U.S.A.,
Calkins 31 (H-NYL 7460!), and Cuba, T, i-'./i
69 (H-NYL 7461!). Calkins 31 was cited as
type by Wirth & Hale (1978), but cannot be
the type as it is not in geographical accordance
with the protologue and also was collected
one year after the description (Staiger 2002).
This material is conspecific with Fissurina
incrustans, as already established by Wirth
& Hale (1978). Wright 267b is a different


Fissurina with similar ascospores, but olive-
brown thallus and very short and broad, widely
open, almost chroodiscoid lirellae with white,
erect margins. This specimen is the same
taxon as Graphina. l i.,.. ... ti., / Fink from
Puerto Rico (Fink 1927), described as having
closed lirellae, but in the type clearly showing
short and wide open lirellae with erect, white
margins. Finally, Wright 69 is yet another
*.p...-.-,. with lirellae of the vlwq.i .i; j .-type,
i.e. roof-shaped and distinctly carbonized.
This material appears to be conspecific with
the new species Fissurina -. i". -, m,\ i ,i/.
(formally described in Lumbsch et al. 2011).
None of the three specimens can conclusively
be designated as lectotype, since they all deviate
from the protologue either in morphological or
label details; on the other hand, it is unlikely
that there are further specimens of this series
which we have not seen, since the collections
were checked hi,-l,-rti in hb. Nylander and at
that time only Cdlla 31 was on loan to Bettina
Staiger. Conceivably, the listing of number 68
in the protologue (based on Nylander 1886)
could be an error for number 69; however,
Wright 69 deviates strongly in its lirellae type
from the description given in Tuckerman
(1888), which on the other hand fits Wright
267b rather well. Tuckerman (1888) obviously
based his description on North American
material (Florida, Ravenel s.n.) and stated
that he had not seen an original description of
Nylander's species. In that case, the validity
of the description can certainly be questioned.
Presently, we apply the name E. nitidescens
in the sense of Wright 267b, pending further
information.
Fissurina aff. elaiocarpa (A. W. Archer) A. W.
Archer.-Figure 24F (Common 7356T, 7425F).
This material can be considered the muriform
counterpart of Fissurina insidiosa. It agrees
%. iI with the description and types ofFissurina
elaiocar)pa (synonym E Ai.,i i;i.,1 Staiger),
except that the ascospores are consistently
I-. Given the taxonomic importance of this
character in the family, the Florida material






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


likely represents an undescribed taxon (the
non-amyloid counterpart ofE elaiocarpa), but
the material is too scanty to allow for a formal
description.
Fissurina humilis (Vain.) Staiger-Figure 24G-
H (Common 72761, 7356M, 7368H).
Fissurina illiterate (R. C. Harris) Lendemer-
Figure 25A (Liicking & Rivas Plata 26618).
The paraphyses in this species are often
thickened at the apex, a feature not observed in
the newly described E analphabetica.
***Fissurina ;.w%,rt- Common & Liicking-
Figure 25B (Common 7276F, 7323C, 73560,
7368J, LI,.. ,l'; & Rivas Plata 26510). See p.
143 for description of this new species.
*Fissurina mexicana (Zahlbr.) Liicking &
Rivas Plata-Figure 25C (Common 7323H,
Common 7418G, Harris & Buck s.n., L .. l; i
& Rivas Plata 26513*, 26514, 26515).
*** Fissurina pseudostromatica Liicking & Rivas
Plata-Figure 25D (Licking & Rivas Plata
26512). See p. 145 for description of this new

*Fissurina radiata Mont.-Figure 25E (Common
7356A 2, 7418E, Licking & Rivas Plata
26619b).
Fissurina rufula (Mont.) Staiger-Figure 25F
(Breuss 2 769*, Seavey & Seavey 10457). The
concept of this species is not clear. We have
not been able to study the type, but according
to Harris (1995), the labia are striate and
ascospores to 24 [im long. The material from
Fakahatchee includes both specimens with
striate and entire labia, and the latter are usually
identified with the name Fissurina subcontexta
(= Graphics ..!i, j, ///. Nyl. in Harris 1995),
supposedly also with shorter ascospores
(15-20 [im). Mason Hale's Index Cards in
US indicate a syntype of Fissurina rufula in
UPS as striate, but with ascospores 13-20 [im
long. Tuckerman (1888) gave the ascospores
of E rufula (the striate form) as 14-18 [im,
and Staiger (2002) included Graphis robustior
MUll. Arg. as synonym of E subcontexta; the
type of that species has robust, partly striate
labia and ascospores about 15-18 um long.


The situation is complicated by Archer's
(2009) synonymization of F subcontexta and
G. robustior with E insidiosa, also suggested
by Staiger (2002), who in addition .u -_.- .1. F
rufula to be the oldest name for this complex.
On the other hand, Harris (1995) maintained
in ..i. .... r, rufula,; -,,,i ..,,.; .,,j i; i f = subcontexta)
separate on account of lirellae morphology and
ascospore size. A further problem is the fact
that, while most forms in this complex have
I- ascospores, there is Fissurina crassilabra
with distinctly amyloid ascospores. As a
consequence of our findings, we distinguish
-p..;-, in this complex as f, 11 , (1) Lirellae
short, erumpent-labiate, thallus uneven-
verrucose, usually yellowish brown= Fissurina
ti..i. .-., ascosporess I-) or E crassilabra
ascosporess I+ violet-blue). Graphics interversa
Nyl. and G. beaumontii Tuck. are synonyms of
either species, but the types have to be checked
for ascospore amyloidity. (2) Lirellae long,
prominent-labiate, entire or becoming striate,
thallus smooth to uneven, usually olive-green,
ascospores I- = Fissurina rufula. Since there
is no apparent difference in ascospore size,
the distinction between entire and striate
forms cannot be maintained, which is also
supported by specimens like the type of G.
robustior, which bears both entire and striate
lirellae. Synonyms of F rufula thus include:
Graphis subcontexta Nyl., G. r-,ii. ,i ii Nyl.,
G. robustior Mull. Arg., and G. rufula var.
comirana Vain.
***Fissurina subcomparimuralis Common &
Liicking-Figure 25G (Common 7276A,
7323A). This new species is formally described
in a separate paper (Lumbsch et al. 2011).
Fissurina subnitidula (Nyl.) Staiger-Figure
25H (Common 7356B, 7368B, 7380A, 7418H,
Licking & Rivas Plata 26511, Seavey &
Seavey l, / ') The concept of this species
in the literature is not accurate. Harris (1995)
and Staiger (2002) described it as having a
non-carbonized excipulum and non-amyloid
ascospores. However, as Tuckerman (1888)
correctly noted, the type shows apical






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


carbonization and this is also seen in all our
collections. The ascospores are persistently
small, not exceeding 15 x 5 pm and usually
about 11-13 jim long, and are weakly to
strongly amyloid. The species thus agrees with
Fissurina ~', /,. ,/i. (Nyl.) Nyl. in having
lirellae of type II according to Staiger (2002),
but the latter has larger ascospores about 13-
23 pm long.
*Fissurina /(ah.Ii 'frapha (Nyl.) Staiger-Figure
26A (Common 7276E, 7413C, 7418A).
***Fissurina tuckermaniana Common &
Liicking-Figure 26B (Common 7266,
7323D). See p. 145 for description of this new

* **Fissurina varieseptata Common & Liicking-
Figure 26C (Common 7413A). See p. 146 for
description of this new *p... i.--
Flakea pipillaii O. E. Erikss.-Figure 26D
(Lay 09-0062). This *-p.... <-. was included in
1i, ::iiiii., (Aptroot et al. 1997) but molecular
phylogenetic analysis suggests that is represents
a separate lineage (Muggia et al. 2009).
*Glyphis atrofusca (Miill. Arg.) Liicking-
Figure 26E Ui, 09-0015). This earlier name
replaces Glyphis montoensis (Archer) Staiger
(Lucking et al. 2009b). The taxon is identical
with GCiilhii, sp. 1271 in Harris (1995:10),
as already noted by Staiger (2002). Graphics
mesoleucodes Nyl. is also conspecific and
antedates Graphina atrofusca Mill. Arg, by
one year, but was published as a nomen nudum
only (Nylander 1886).
Glyphis cicatricosa Ach.-Figure 26F-G
(Beeching 7600, 7685, Buck 54438, Common
7259C, 7277B, 7292E, H. ..i//,,. i 10-19, Lay
09-0073, Mercado-Diaz 412, Liicking & Rivas
Plata 26619*, Seavey & Seavey 10067). The
abundant material includes both forms with
stroma bearing numerous rounded to slightly
elongate lirellae, as well as forms with stroma
bearing a single, radiately branched lirella. The
stroma are uniform within a given individual,
but thalli with intermediate forms have also
been found, although they are rare. Possibly
two distinct taxa are involved here and DNA


studies will be needed to solve the issue.
fhj',iu' scyphulifera (Ach.) Staiger-Figure 26H
(Common 7292D). In the checklist (Esslinger
2010) as Gyrosiomum scyphuliferum (Ach.)
Nyl.
Graphis cf. acharii Fee-Figure 27A (IlAu/r:
& Rivas Plata 26521*). The material is
unfortunately sterile but agrees in the thallus
and lirellae morphology as well as the clear
hymenium with Graphis acharii, which has
muriform ascospores. Other likely alternatives
would be the less common Graphis ai, _nu.ri.ii,n
Eschw. ascosporess transversely septate) and
Graphics vestitoides (Fink) Staiger ascosporess
terminally muriform). The genus Graphis was
keyed out with over 300 species world-wide by
Lucking et al. (2009b); molecular phylogenetic
analysis -u--i- -1 that it actually includes two
ItI.-;rh.l related lineages (Rivas Plata et al.
2011).
Graphis anfractuosa Eschw.-Figure 27B
(P.. chli-r 7644, Lay 09-0070, Seavey &
Seavey 10319). Graphis anfractuosa forms
part of a misunderstood complex of species
with completely carbonized excipulum and
inspersed hymenium (Table 5). Specimens
with small ascospores and norstictic acid
represent either G. desquamescens (disc
concealed) or G. aperiens (disc exposed),
whereas specimens with larger ascospores and
lacking secondary substances are either G.
anfractuosa ascosporess 30-40 ptm long) or G.
cupei ascosporess 40-60 ptm long).
Graphis aperiens Miill. Arg.-Figure 27C
(Common 7425D).
***Graphis appendiculata Common &
Liicking-Figure 27 D-E (Common 7313A,
7368L, 7-:..'.'T!, 7418K, Lay 09-0024, 09-0068,
Licking & Rivas Plata 26657*, 26659*,
26660*, 26668*, Mercado-Diaz 411). See p.
147 for description of this new species.
*Graphis argentata Liicking & Umafia.-Figure
27F (Breuss 28825*).
*Graphis assimilis Nyl.-Figure 27G (Common
72590, 7276K, 73461).
Graphics caesiella Vain.-Figure 27H (Buck







LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


Table 5. Simplified keytable to identify species of Graphis cited in this work. Septa = number of ascospore septa: muri
= muriform; excipulum = excipulum carbonization; inspersion = hymenium inspersion; labia = labia striation; chemistry =
secondary chemistry; lirellae = lirellae emergence; margin = thalline margin of lirellae; other = other characters: exposed =
Ii .. .1 disc; green = green thallus; pruinose = pruinose labia; verrucose = verrucose i.l.in size = ascospore size in ttm
(typical upper range for length and size; the typical lower ii.n is about 30% less).


Species of Septa Excipulum Inspersion Labia ( icli,;i. Lirellae Margin Other Size
Graphis


G. oshioi

G ..... : .. ...

G. appendiculata

G. caribica

G. caesiella

G . .. ..

G. leptocarpa

G. handelii

G. sauroidea

G. assimilis

G. caesiocarpa

G. conferta

G. oxyclada

G. stellata

G. rimulosa

G. longula

G. desquamescens

G. aperiens

G. anfiractuosa

G. cupei

G. disserpens

G. xylophaga

G. acharii

G. subflexibilis

G. arena.
G. argentata


5-9

7-9

9-13

9-15

5-9

5-7

7-11

7-9

7-15

7-11

7-11

7-11

7-11

5-7

7-11

11-17

5-9

5-9

7-11

13-17

muri

muri

muri

term

muri

muri


absent

apical

Si .

apical

lateral

lateral

lateral

lateral

complete

complete

complete

complete

complete

complete

complete

complete

complete

complete

complete

complete

apical

lateral

complete

complete

complete

complete


ins

ins


ins]

ins]

ins]

ins]


ins]

ins]

ins]


- entire norstictic

- entire -

- striate -

- striate -

- entire norstictic

- striate lichexanth

versed entire stictic

versed entire norstictic

- entire lichexanth

- entire norstictic

- entire norstictic

- entire -

-- entire -

- entire -

- striate -

- striate -

persed entire norstictic

3ersed entire norstictic

versed entire -

persed entire -

- striate -

-- entire -

- striate -

persed striate -

3ersed striate -

3ersed striate -


prominent

emrmpent



prominent

emrmpent

ennnpent

erumpent

emmpent

prominent



erumpent

sessile

prominent

prominent

erumpent



erumpent

emrmpent

prominent

prominent

emrmpent

enunpent

prominent

prominent

prominent

prominent


complete

lateral

comp thin

absent

lateral

absent

lateral

lateral

basal

lateral

comp thin

absent

lateral thick

lateral

absent

lateral

basal

lateral

absent

basal

lateral

lateral

comp thin

comp thin

comp thin

comp thin


S 30 x 7

pruinose 35 x 8

55 x 12

green 60 x 12

pruinose 35 x 7

30 x 8

35x 7

exposed 40 x 8

50 x 12

40 x 8

pruinose 35 x 8

40 x 8

40 x 8

verrucose 25 x 7

50 x 12

65 x 12

S 25 x 7

-: -., ,1 30 x 8

35 x 8

55 x 8

40 x 20

-- 80 x 25

120 x 20

120 x 18

60 x 12

100 x 20


1






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


54439, Common 7413B, 7425B, Liicking &
Rivas Plata 26524*, Seavey & Seavey 10362).
*Graphis caesiocarpa Redinger-Figure 28A
(Breuss 28718*).
*Graphis caribica Liicking-Figure 28B
(Common 7356G). This species was already
discussed in Licking et al. (2008) for Costa
Rica and is formally described in a separate
paper (Lumbsch et al. 2011).
*G,.rap'i, conferta Zenker.-Figure 28C
(E, -, hi u- 7602, Lay 09-0016, Mercado-Diaz
441).
*Graphis cupei Vain. ex Luicking-Figure 28D
(Common 7380H, Licking & Rivas Plata
26526*).
Graphics ,: \i,,,t,,.:,,.,, (Fee) Zahlbr.-Figure
28E (Common 73680, Lay 09-0064, Seavey
& Seavey 10377*). We are uncertain about
the correct application of the name Ci. ;hi..
..,,.,, ...... The Florida material has
very small ascospores to 25 pm and the outer
wall of the end cells is amyloid, a feature
rarely observed in other Graphis species. The
lectotype has ascospores 25-35 pm long and
other material from the Neotropics fits this
size range; the end cell walls are usually not
amyloid. It is possible that different species
are involved here, but we want to await more
detailed studies on ascospore morphology and
chemistry in a wide range of G,.., :/i. species
before using this character formally.
*(G,.,Y;h; disserpens Nyl.-Figure 28F-G (Lay
09-0224, kicking & Rivas Plata 26656*).
*(Gr,'q;h handelii Zahlbr.-Figure 28H (Common
7292G).
Graphics leptocarpa F6e-(Common 7346E).
*Graphis lm upu Kremp.-Figure 29A (Common
7380B, L i;,, I ir,- & Rivas Plata 26527).
Graphics hin ', p R. C. Harris-Figure 29B (Layo
09-0065).
*Graphis oshioi M. Nakan.-Figure 29C ':;';.**..
28706* Common 7 7'C, 7313H, Lay 09-
0076, Lciiking & Rivas Plata 26612). This
species would be classified as a Hemithecium
in the concept of Staiger (2002), because
of the absence of excipulum carbonization.
However, molecular data show that species of


Hemithecium are nested within Graphis, and
the differences in i- ..,_iukli1 carbonization
are merely gradual (Rivas Plata et al. 2011).
Graphics oshioi is similar to the widespread
tropical Graphis ;iyi.pi. ,a.' Fee and differs
mainly in the presence of norstictic acid.
Graphis oxyclada Miill. Arg.-Figure 29D
(Common 7 -.1 I/ 7380J, H, ..... s.n., Lay 09-
0017C, 09-0173, Lendemer 15683, Licking &
Rivas Plata 26528a).
*Graphis pseudocinerea Liicking-Figure 29E-
G (E... lc./,in 7630, Breuss _''\, 7*, Lendemer
15547, Liicking & Rivas Plata 26533*, 2653 7*,
Mercado-Diaz 392, Seavey & Seavey 10337).
Graphis rimulosa (Mont.) Trevis.-Figure 29H
(I. l/,;;: & Rivas Plata 2653-i).
*Graphis sauroidea Leight.-Figure 30A (Breuss
".n '.', Lendemer 15546).
*Graphis stellata M. CAceres & Liicking-
Figure 30C-F (Breuss .'"-- ." Lay 09-0067,
Seavey & Seavey 10321). This find represents
and interesting disjunction, as the species was
previously only known from northeastern
Brazil (Caceres 2007). The Florida material
agrees well with the type, except that the
ascospores are 5-septate rather than 7-septate
and at the lower side of the size range (20 tpm
versus 20-30 tm). As in other cases, we expect
this species to be also found in the Caribbean
and/or parts of Central America.
*Graphis subflexibilis Liicking & Chaves.-
Figure 30B (Common 7356H, 7368E, 7-.'.' ,Gt).
*Graphis xanthospora Miill. Arg.-Figure 30G
(Breuss ','._*, Lendemer 15537, Lficking &
Rivas Plata 26535*).
Graphics xylophaga (R. C. Harris) Lendemer-
Figure 30H (Crane ILLS 60442, ',. ,vey &
Seavey 10359).
CGi, id, iid ii appendiculatum Liicking &
Lendemer-Figure 31A-B (Sanders P1:~-4.4,
10504.6a).
Gyalectidium catenulatum (Cavalc. & A. A.
Silva) L. I. Ferraro, Liicking & Serus.-
Figure 31C (Sanders It.L-1.8).
CGi, ,I ,idimin flui h'ii,'c Safranek & Liicking-
Figure 31D-E (Sanders 0l-1'ii 3, 10521.- )
QGlc iidifhiiim imperfectum VWzda-Figure 31F






LUCKING ET AL.: Lichens of Fakahatchee t.I i Preserve State Park


(Liicking & R',,.. Plata 26808b, Mercado-
Diaz 419d, Sanders 10504.7).
*(, ,Il, A1- FInhn niLa, Herrera-Campos &
Liicking-Figure 31G (Sanders 1],,l i. 10).
Gyalectidium. aff. yahriae W. R. Buck & Serus.-
Figure 31H (Sanders 10521.2, 10521.3). This
material resembles Gyalectidiumyahriae in the
more or less radially symmetrical hyphophores
with a crater-like base and the irregularly
incised appendages, but differs clearly in
that the diahyphae form a compact mass
embedded in a gelatinous matrix. Gyalectidium
denticulatum Licking is somewhat similar but
has a verrucose instead of smooth thallus and
the hyphophores have a thinner margin and
more delicate appendages. Unfortunately, the
material is too scanty to allow for a formal
description.
Haematomma fl-'vrumii Hillm.-Figure 32A
(_.-,_./,, ,, .!i ) ni).
Haematomma guyanense Kalb & Staiger-
Figure 32B (Harris & Buck s.n., Mercado-


Diaz ,i "*' ).
Haematommapersoonii (Fee) A. Massal.-
32C (Seavey & Seavey 10070).
Hf'cll;ni bahiana (Malme) Sheard.-
32D-E (Harris & Buck s.n.).
Hf 'cll;n curatellae (Malme) Marbach-
32F (Lendemer 15542A, Seavey &
10615).


-Figure

-Figure

-Figure
Seavey


Haftlia pl,'ir-ira (\lidt R) Marbach-Figure
32G (Breuss 28936*).
***riiI'.:, rPhafflwiiLdn,, Common & Liicking-
Figure 32H (Common 7410C). This new genus
and species is described by Licking et al.
(2011). The genus is characterizedby combining
a Cir. ,. hi -like whitish thallus and carbonized,
labiate lirellae with a P./ ...',"I- t.y''.-like in-
spersed hymenium and gray-brown, Phaeogra-
i'lni. ype ascospores (short with rounded
ends and thick walls compared to longer with
tapering ends and thinner walls in GC..,,':/;.. It
is phylogenetically distinct from both Ci. ,1.hi,
and P!,,..' '1,7 ,i; s.str., although more closely
related to the latter.
***Heiomasia w.',!ri-, In,, M. P. Nelsen &


Liicking-Figure 33A-B (Beeching s.n.,
Licking & R,.... Plata 26850, Nelsen 4076).
This new genus, with two new species, was
described by Nelsen et al. (2010). It is a genus of
two sterile taxa based on L-',i i ,i. i. v sipmanii
Aptroot et al. from the Palaeotropics (Aptroot
et al. 2009). The North American species
has sausage-shaped isidia and an unusual
chemistry of 5-hydroxy-4-O-demethylnotatic
acid and 4-O-demethylnotatic acid.
f -ipiriatimnM antillarum (Vain.) Aptroot,
Liicking & G. Thor-Figure 33C-D (Nelsen
?7).
*H:'DnDl!,n echinatum Aptroot, Liicking
& Will-Wolf.-Figure 33E (Lay 09-0008,
Licking & Rivas Plata 2<'. '.1*).
fir,'wdrrfaii I rubrocinctum (Ehrenb.) Aptroot,
Liicking & G. Thor-Figure 33F-G (Buck
5/78, h,. mi .., 10555, Lay 09-0075,
Lendemer 15624, Licking & Rivas Plata
26707, 26708*, Nelsen , or., Seavey & Seavey
10725).
f/,'irpflalriniu rubroechinatum A. Frisch & G.
Thor-Figure 33H (Licking & Rivas Plata
26682). This species was recently established
based on material from southern Florida
(Frisch et al. 2010).
Heterodermia albicans (Pers.) Swinscow &
Krog.-Figure 34A (:;'.,, .. 28763*, Lay 09-
0077).
Heterodermia pseudospeciosa (Kurok.) W. L.
Culb.-Figure 34B (Breuss ;"' ,' ).
Laurera megasperma (Mont.) Riddle-Figure
34C-D (P.: i,,!;i 7: ', Crane ILLS 60461,
Lay 09-0080, Lendemer 15580, T i;, .i, &
Rivas Plata 26709, 26710a*, Nelsen 4015,
Seavey & Seavey l, ').
*Lecanora achroa Nyl.-Figure 34E (Seavey &
Seavey 101O i:)
Lecanora achroides Vain.-Figure 34F (Seavey &
Seavey 10624*).
Lecanora ,IhnlJhnn! Nyl.-Figure 34G (Licking
& Rivas Plata 26639).
Lecanora argentata (Ach.) Malme-Figure 34H
(Lay 09-0136, Liicking & Rivas Plata 2663 ',',
26636*, Seavey & Seavey 10631).






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


Lecanora 'a,' ,wuh,'h/l, ssp. glaucomodes (Nyl.)
Imshaug & Brodo-Figure 35A (Seavey &
Seavey 10077*).
*Lecanora .tp.-i;,,n Stizenb.-(Buck 5 / 1r-,
Lendemer 15577).
Lecanora fl4oI.;,hIl Lumbsch-Figure 35B-C
(. hi1u. 7675, Buck 54 ." /, Lay 09-0083,
Lendemer 15528, Seavey & Seavey 1,. i.';'*).
Lecanora y hi, -iarpi (Tuck.) Brodo-Figure 35D
(Lay 09-01.. ')..
Lecanora leprosa Fee-Figure 35E y:.';..
"'-1.:';*, Lendemer 15550, Seavey & Seavey
10079*).
Lecanora /,.' 1;.'b',' Lumbsch-Figure
35F (Ti;, kiX; & Rivas Plata 26642, Seavey &
Seavey 10081).
Lecanora strobilina (Spreng.) Kieffer-Figure
35G (Licking & Rivas Plata 26637*, 26638*,
Seavey & Seavey l o; "').
Lecanora spec.-(Mercado-Diaz 399). This
material is characterized by beige apothecia
with thin margin, much as in Lecanora helva
Stizenb., but has numerous small crystals
in the amphithecium that dissolve in K and
lacks crystals in the epithecium. The only
species matching this apothecial anatomy is
L. epibryon (Ach.) Ach., which has apothecia
with much darker discs and thicker margin.
Leiorreuma exaltatum (Mont. & Bosch) Staiger-
Figure 35H (Common 73551, 7424A).
Leiorreuma cxjr,\ wh# (Fik) Lendemer-
(Beeching 7637).
Leiorreuman sericeum (Eschw.) Staiger-Figure
36A (Beeching 7625, Common 7313F, 7355E,
742h /G, Lendemer 15557, Seavey & Seavey
1i '.. 3).
Leptogium i'vau '\,',n\ (Rabenh.) Kirb.-
(Beeching 7586, Lay 09-0086, Lendemer
15614, Seavey & Seavey 1" /-,).
Leptogium denticulatum Nyl.-(H ,. /, in.,
10557, Seavey & Seavey lo ' ").
Leptogium i',ti% d.lhi, (Riddle) Sierk.-
(B.-, hi 7638, H .. iilm ...a J 1 /2).
Leptogium marginellum (Sw.) Gray-(,v, '.' iw:;
7721, Buck 54450, Crane ILLS ,., *',
Hodkinson l1. _i2, Lay 09-0089, Lendemer


15631, Seavey & Seavey el 191*).
Leptogium microstictum Vain.-(Seavey &
Seavey 10493).
Leptogium milligranum Sierk.-(Mercado-Diaz
462, Seavey & ',. .,ey 1022 i).
Letrouitia domingensis (Pers.) Hafellner &
Bellem.-Figure 36B (Beeching 7587,
Common 7379, H. ,J,;,r.. vi 1-_.';, Lay 09-
0093, Licking & Rivas Plata 26717*, 26718,
Nelsen 4u-'.3b, Seavey & Seavey 10225*).
Letrouitia vulpina (Tuck.) Hafellner & Bellem.-
Figure 36C-D (Crane ILLS ,o44.'", Lay 09-
1~ 'v I, Lendemer 15621, Licking & Rivas Plata
26723*, Mercado-Diaz 390, Nelsen -II::...
Seavey & ',. .,vey 10539*).
*Leucodecton conpunctellum (Nyl.) A.
Frisch.-Figure 36E (Lucking & Rivas
Plata 26566b, Seavey & Seavey lai _'_). The
genus Leucodecton is treated with a key to all
accepted species by Rivas Plata et al. (2010).
Leucodecton glaucescens (Nyl.) A. Frisch.-
Figure 36F-G (E...... air 7588, L.A..,,/: &
Rivas /!'/. a 26538, 26539*, Nelsen 4182,
Seavey & Seavey 10230).
Leucodecton occultum (Eschw.) A. Frisch.-
Figure 36H (Common 7321B).
Lithothelium microsporum R. C. Harris-(Harris
& Buck s.n.).
Malmidea furfurosa (Tuck. ex Nyl.) Kalb &
Liicking-Figure 37A (Lay 09-0098, Seavey
& ',.iey 10232). Species of this genus
were until recently treated under the name
Malcolmiella (Caceres 2007), but molecular
phylogenetic analysis showed that the type
species of \ i ,'.. :la. /,, is unrelated to the
remaining taxa placed in that genus, and the
new genus Malmidea was erected for this group
(Kalb et al. 2011). Most species can be keyed
out using the treatment under \ ';,, ,h:i. I/,, in
Caceres (2007) plus the key to Thai species
of Malmidea provided by Kalb et al. (2011).
Manmidea fli,,' ....1 is currently listed as
Lecidea jJii dI ,..-: Tuck. ex Nyl. in the North
American lichen checklist (Esslinger 2010).
*Malmidea fuscella (Miill. Arg.) Kalb &
Liicking-Figure 37B (Breuss -.. 711*).






LUCKING ET AL.: Lichens of Fakahatchee i.i i Preserve State Park


llliniJf, granifera (Ach.) Kalb, Rivas Plata
& Lumbsch-Figure 37C (Beeching 7632,
Breuss 28711*, Buck 5 ,, w, Crane ILLS
i.o i 78, Licking & Rivas Plata 26725*, Seavey
& Seavey 10314).
*Malmidea gyalectoides (Vain.) Kalb &
Liicking-Figure 37D (Licking & Rivas Plata
26727, 26728*, Nelsen 1, 17).
*raLdmnril.- leptoloma (Miill. Arg.) Kalb &
Liicking-Figure 37E (Lendemer 15564,
Liicking & Rivas Plata 26730).
*Malmidea piperis (Spreng.) Kalb, Rivas Plata
& Lumbsch-Figure 37F (Lay 09-0090).
*Malmidea rhodopis (Tuck.) Kalb, Rivas Plata
& Lumbsch--(/;', : .. .*''::7*).
*Malmidea variabilis Kalb-Figure 37G
(. .*, hii; 7585). This taxon was recently
described from Thailand (Kalb et al. 2011),
being somewhat similar to M. i.~,; s,., but
differing chiefly in the papillose apothecial
margin. Similar collections have been reported
from Brazil (Caceres 2007) but were left un-
named. Our identification is tentative because
in the Neotropics apparently two species with
slightly different chemistry are involved which
need further study.
*rLdnarii,,Li vinosa (Eschw.) Kalb, Rivas Plata
& Lumbsch-Figure 37H (Breuss 28658*,
28690*, Lay 09-0099, Liicking & Rivas Plata
26731).
1L',plq',,lra porphyritis (Tuck.) R. C. Harris-
Figure 38A-B (Harris & Buck s.n, Hodges
s.n.).
M..;iJnir,,',mi spec.-(Lay 09-0227, Licking
& Rivas Plata 2;*'.-'). This material is very
characteristic because of its comparatively
large (35-50 x 10-14 ipm), distally pointed,
ornamented ascospores with markedly sub-
median septum. The thallus is endoperiderm-
al and white and patchily UV+ yellow
(lichexanthone). The perithecia are in r ,miit nt
black, and slightly irregular although with
more or less apical ostiole. It does not key out
to any of the species ofAnisomeridium s.lat. in
Harris (1995); because of its large, ornamented
ascospores, it seems to belong iii 1 / ."' 'I ,


sensu Aptroot (1991), but none of the species
currently accepted in that genus comes close.
The recently described M. lateralis Aptroot has
perithecia with markedly excentric ostiole and
much broader ascospores with the distal end
rounded (Aptroot et al. 2008).
Micareaprasina Fr.-Figure 38C (Lay 09 ,r.'. ".
Micarea spec.-Figure 38D-G (Lay 09-0051).
This unidentified material bears numerous
apothecia and tubular pycnidia and probably
represents an undescribed species.
MyJ y 38H (Buck 5 / i71, Lay 09-0097).
MlrJ Hawksw.-Figure 39A (Licking & Rivas
Plata 2r..- ":,. *).
i ,,,imiiliJ,,hlit subfallens (Miill. Arg.) D.
Hawksw.-Figure 39B (&; ,,i:. & R;j../.
/'!,I / 26772*, Seavey & ',. .,ey 10821).
M Hawksw.-Figure 39C-D (Licking & Rivas
Plata 26597*, 26599, 26733*, Mercado-Diaz
S/ i). The species resembles i .i ., ; ,,,ii,
especially as the ascospores are often pale to
hardly pigmented at all. I ,., :' i. ,rh/ i. (at
least the three species sequenced to date) has
recentlybeen showntobelonginTrypetheliaceae
(Nelsen et al. 2009), which is supported by the
thin, distinct, anastomosing, loosely net-like
interascal hyphae. ir. i. u.. ..,in/ has a similar
type of hamathecium and also belongs in
Dothideomycetes, but in a separate clade distant
from Trypetheliaceae (Nelsen et al. 2009). If
ascospore pigmentation is variable in both
Anisomeridium and \ i, ii-. i -ili. !.,, some
S....i.-<-, currently placed in Anisomeridium
could belong in 3 !. n, viii1i i.,, and vice
versa. The situation is complicated by similar
looking species of fli :ili (another clade
in the Dothideomycetes) with branched and
anastomosing interascal hyphae (Nelsen et al.
2009).
Mycoporum buckii R. C. Harris-(Harris & Buck
s.n.).
Mycoporum eschweileri (Miill. Arg.) R. C.
Harris-(Beeching 7581, Buck 54403).






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


Mycoporum lacteum (Ach.) R. C. Harris-Figure
39E-F (Liicking & Rivas Plata 26732).
Mycoporum \pE~nI.lihn Nyl.-Figure 39G-H
(Liicking & Rivas Plata 26613).
1tyri,ar-irv erodens R. C. Harris-Figure 40A-
B (P, ._-, /;1. 7611, Lendemer 15687A, Lay 09-
0102, Liicking & Rivas Plata 26541, Mercado-
Diaz 436, Nelsen 4183, Seavey & Seavey
10811*). Thalli with the coarse, somewhat
diffuse soralia typical of Myriotrema erodens
were found quite abundantly at Fakahatchee.
Molecular analysis of some sorediate specimens
(Rivas Plata et al., unpubl. data) showed that
they are very similar to apotheciate specimens
identified as O, .'l//;/. ,i / auberianoides (see
below). The latter agrees in most features
with fertile Myriotrema erodens except for the
distinctly smaller ascospores and more open
apothecia with irregular columella structures.
Myriotrema erodens is based on material from
Cuba and so far only the type and an additional
specimen from Panama (Harris 2' '- T) have
been found with apothecia. Coincidentally,
a sorediate species with small ascospores,
C i ,'ii li, ;., .,.., .h7;.j, ., Kalb, was recently
described from the Neotropics (Kalb 2009),
which corresponds to forms with apothecia of
the 0. auberianoides type and soralia of the
Myriotrema erodens type. Kalb (2009) indeed
suggested O. ,..,,.,.il,_... to be the sorediate
counterpart of 0. auberianoides. In the present
material, we have not found any specimen
with apothecia and soralia on the same thallus;
we therefore used Ocellularia auberianoides
for apotheciate material and Myriotrema
erodens for sorediate specimens, but molecular
analysis of a large number of -i iir, r,... could
well reveal the presence of up to three taxa,
Myriotrema erodens s.str., 0. auberianoides,
and 0. -- '..h. i ._,,. It is also possible that all
sorediate North American material hitherto
identified as Myriotrema erodens belongs
to O. ,'i' ,/,, ., sorediigera or even that that
0. auberianoides and 0. ..-,./. ''..-..., are
conspecific, in which case only one taxon
would be recognized.


.li'ih-,r;',-' peninsula R. C. Harris-Figure
40C (Licking & Rivas Plata 26542*, Nelsen
4181, Seavey & Seavey Io'. /7).
*_/i riiit.i,, JtB'yii, iq,,.:1].,ll,, (Nyl.) Hale- Figure
40D (Beeching 7666, Common 7359A, 7371B,
7377B, Lay 09-0i' _,, Licking & Rivas Plata
26544*, Seavey & Seavey 10133).
Nadvornikia hawaiiensis (Tuck.) Tibell-Figure
40E (Breuss '. .'.'I .
Nadvornikia 'ir,:.ianott R. C. Harris-Figure 40F
(Lendemer 155 s t).
* 0.:,llul trin auberianoides (Nyl.) Miill.
Arg.-Figure 40G (Lay 09-o"' 7, Liicking
& Rivas Plata 26548*). This taxon has been
synonymized with 0. bonplandii (Fee) Mill.
Arg., but differs in the larger apothecia with
irregular-reticulate pseudocolumella; both are
also genetically different. Ocellularia auberi-
anoides closely resembles 0. obturascens but
can be readily distinguished by the hyaline,
transversely septate ascospores (brown and
muriform in 0. obturascens).
*0'4.-lIIin'i, obturascens (Nyl.) Hale-Figure
40H (,-., /./ 7720, Lay 09-0103, Lendemer
15551, 15627, f i;, Ni;u & Rivas Plata 26552*,
2(:-. /', 26555*, Mercado-Diaz 381, Seavey &
Seavey 10136*). The material ofthistaxon from
southeastern United States is usually identified
as Myriotrema bahianum (Ach.) Hale, recently
recombined as Ou. ',!i, i..t bahiana (Ach.) A.
Frisch (Frisch & Kalb 2006). However, that
species lacks a columella or very rarely has
a rudimentary pseudocolumella, whereas the
Florida material is without exception distinctly
columellate. The name 0, '77I,'. i. obturascens
is available for the columellate taxon; this name
is listed as synonym of 0. bahiana in the North
American lichen checklist (Esslinger 2010) but
should be restored and 0. bahiana be deleted.
Ochrolechia ,!fri:,u-ni, Vain.-Figure 41A (Seavey
& Seavey 10752*).
Ochrolechia antillarum Brodo-(Lendemer
15642).
Opegrapha astraea Tuck.-Figure 41B-C
(Licking & Rivas Plata 26734*, Mercado-
Diaz / f/, Seavey & Seavey 10741).






LUCKING ET AL.: Lichens of Fakahatchee ti. i Preserve State Park


Opegrapha atra Pers.-Figure 41D (Licking &
Rivas Plata 2673 7a*).
Opegrapha candida Miill. Arg.-Figure 41E
(Seavey & Seavey 10628*).
Opegrapha cypress R. C. Harris-Figure 41F
(Breuss 28715*, Lendemer 15588, Seavey &
Seavey 10791).
Opegrapha longissima Miill. Arg.-Figure 41G
(Harris & Buck s.n., Licking & Rivas Plata
26736, 26815).
Opegrapha spec.-Figure 41H (Mercado-Diaz
450). This material is the same as Britton 652
cited in Harris (1995: 7). It is characterized
by robust, sessile lirellae with basally closed
excipulum and slit-like disc, inspersed
hymenium, and 7-septate ascospores 40-45
x 5-7 ipm in size with the two median cells
slightly enlarged. Most similar appear to be
0. varia Pers., with smaller, usually 5-septate
ascospores and non-inspersed hymenium, and
the paleotropical 0. ;,.,, .~.-.i.. Ertz, with non-
inspersed hymenium, more delicate lirellae,
and smaller, 6-septate ascospores.
Parmeliella '[ltylph' l~n (Vain.) P. M. Jorg.-
Figure 42A-B (Licking & Rl',,-. Plata 26738).
Parmotrema austrosinense (Zahlbr.) Hale-
(H'a.- hi. 7647).
Parmotrema crinitum (Ach.) M. Choisy-(Seavey
& Seavey 1. I /).
Parmotrema c(r tif,_.,rr (Taylor) Hale-
(P. .;', hi; 7714, Hodkinson le'; i I, Lendemer
15644, Seavey & Seavey 10639).
Parmotrema dilatatum (Vain.) Hale-(Lendemer
15691, Seavey & Seavey 10156*).
Parmotrema endosulphureum (Hillm.) Hale-
(Lendemer 15555, Seavey & Seavey 10559*).
Parmotrema gardneri (C. W. Dodge) Serus.-
(Lendemer 15672).
Parmotrema hypoleucinum (Steiner) Hale-
(Breuss 2.'-.' 3* Seavey & Seavey 105 i,).
Parmotrema perforatum (Wulfen) A. Massal.-
(Crane ILLS ,'.j 13).
Parmotrema sulphuratum (Nees & Flot.) Hale-
(Beeching 7670, Lay 09-0105, Lendemer
1565 i).
Parmotrema tinctorum (Delise ex Nyl.) Hale-


(Crane ILLS 60446, Seavey & Seavey 10158*).
Pertusaria commutata Mill. Arg.-(Lendemer
15637).
Pertusaria -pl'i\ ,nth, R. C. Harris-Figure 42C-
D (Liicking & Rivas Plata 26666c).
Pertusaria t, .tdi; R. C. Harris-Figure 42E-F
(Liicking & Rivas Plata 26650*). The whitish,
sterile sorediate thallus containing stictic acid
and coronatone characterize this species.
Pertusaria aff. expolita R. C. Harris-Figure
42G-H (Liicking & Rivas Plata 2'.; i') This
material agrees with P. 'i '. ,i in the sorediate
thallus and chemistry; however, the soralia
are much smaller and discrete, resembling
goniocystangia, and the thallus is greenish
rather than whitish.
Pertusaria fl.'uiidar, Dibben-(Harris & Buck
s.n.).
Pertusaria I, '/I.Ira DC.-Figure 43A-B
(Licking & Rivas Plata 26623c, Seavey &
Seavey 101-i7*).
*Pertusaria .rtrl h,,.irlj,,t. Dibben-Figure
43C ( :'.*; ..; '"-,' ;*, Liicking & Rivas Plata
26; :-,, 26645*).
Pertusaria sinusmexicani Dibben-Figure 43D
(Lendemer 15682, Liicking & Ri,, Plata
26652*, 2665 i)
Pertusaria tetrathalamia (Fee) Nyl.-Figure 43E
(Harris & Buck s.n., Seavey & Seavey 13332*).
Pertusaria texana Miill. Arg.-Figure 43F
(Lcicking & Rivas Plata 26643, Seavey &
Seavey 10215).
Pertusaria virensica R. C. Harris-Figure 43G
(Lay 09-0141).
Pertusaria .Nv, i J,?'k Miill. Arg.-Figure 43H
(Lendemer 15568, Liicking & RAi, Plata
2,'. i7*, 26651*, Seavey & Seavey 10218).
Ph,~,~.e~o qr1pii l asteroides (Fink) Lendemer-
Figure 44A (Beeching 7665, Lay 09-0145,
Licking & Rivas Plata 26739*, Seavey &
Seavey 10216). Species of PhI .. --,.hi.. found
in Florida are summarized in Table 6.
Phrai"irapihi brasiliensis (A. Massal.) Kalb &
Matthes-Leicht.-Figure 44B-C (Common
7245A, 7355M, 742-i/C, Lay 09-01/ .j Seavey
& Seavey 10266). Some of the specimens







BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. ',, 1,


Table 6. Keytable to identify species of Phaeographis and allied genera (Leiorreuma, / .'. Sarcographa) cited in this
work or otherwise reported from Florida. Septa = number of ascospore septa: muri = muriform; insp = hymenium inspersion;
chem = secondary chemistry: pigm = quinone i;Li.mic (K+ green), lichx= i..-Pi.- cii,-,., exci= excipulum carbonization;
hypo = hypothecium (or basal I ,. of excipulum) carbonization; i ,. I I = lirellae hi. disc = disc ,. 1" m1'. i L,, ; ., other =
other characters: ecorticate = ecorticate thallus, lobulate = lobulate lirellae margin, orange = orange hymenium, striate = striate
labia; size = ascospore size in Inm (typical upper range for length and size; the typical lower range is about 30% less).

Species Septa Insp Chem Exci Hypo Lirellae Disc Other Size


Phaeographis
P haematites
P inconspicua
P subfulgurata
P. major
P brasiliensis
P intricans
P schizoloma
P.flavescens
P. delicatirla
P aff. schizoloma
P punctiformis
P. nylanderi
P dendritica
f .i !ltt I
P inusta
P erumpens
P lobata
P leiogrammodes
P asteroides
P multicolor
P scalpturata
P aff. scalpturata

Platygramme
P praestans
P pachnodes
P caesiopruinosa
P aff. caesiopruinosa


Leiorreuma
L. sericeum
L. exaltatum
L. explicans

Sarcographa
S. labyrinthica
S. tricosa


5-9
3
3
5
3
5
3-5
5
3
5-7
5
3-5
5-11
3
3-5
5-7
5-11
muri
muri
muri
muri
muri



9-17
muri
muri
muri



3
5-9
muri


- pigm
- norst
- norst
- norst
- norst
- norst
- stictic
- stictic
- stictic


lichx
norst
norst


norst


insp norst
insp -
insp -


elongate-stellate
elongate
elongate
elongate
pseudostromatic
pseudostromatic
elongate
pseudostromatic
-stellate
elongate
thin thin round-oval
oval-elongate
thin thin elongate-stellate
elongate
thin thin elongate
thin thin elongate
thick thin round
elongate-stellate
-stellate
thin elongate
- elongate
thin thin elongate


- thick
norst thick
- thick
- thick


thick
thick
thick


3 insp stictic thin thick
3 insp thin thin


elongate
elongate
elongate
elongate



stellate
elongate
elongate



stromatic
stellate


dark red
grey-pminose
grey-pruminose
grey-pminose
grey-pruminose
grey-pminose
brown (p iII i ., )
grey-pminose
brown
brown (pruinose)
brown-black
brown-black
brown-black
brown-black
brown
brown-black
brown (jiI i
grey-pminose
grey-pminose
brown-black
brown (jiI i
brown (pruinose)



bluish pruinose
white-pruinose
bluish pruinose
brown (jiI i ;



brown-black
brown-black
brown-black



grey-pruinose
(pruinose)


30 x 8
13 x 6
lobulate 20 x 9
22 x 8
17 x 6
22 x 6
striate 25 x 10
25 x 6
25 x 9
striate 30 x 10
25 x 8
20 x 6
ecorticate 40 x 10
lobulate 18 x 8
25 x 7
lobulate 30 x 8
lobulate 40 x 10
20 x 8
35 x 12
orange 35 x 10
100 x 30
100 x 20



striate 70 x 13
35 x 10
90 x 20
90 x 20


18 x 7
30 x 8
28 x 11



20x6
20 x 6






LUCKING ET AL.: Lichens of Fakahatchee t. i Preserve State Park


exhibited a negative K-reaction of thallus and
lirellae, although TLC showed the presence of
norstictic acid. Apparently, either the substance
is concentrated in certain parts of the thallus or
occurs in concentrations too low to be reliably
detected by 10% KOH solution. The specimen
supposedly devoid of lichen substances,
Harris 23697 in Harris (1995), was tested
TLC negative and might represent a separate
taxon. Specimens with stictic acid belong to P.
flavescens.
***Phaeographis delicatula Common &
Liicking-Figure 44D (Common 7355J,
7367C, 7381D, Lay 09-0151). See p. 148 for
description of this new species.
Pha,,,-'jri',hl, erumpens (Nyl.) Mill. Arg.-
Figure 44E (Breuss 28734*, Common 7277A,
7367E).
*Phaeographis flavescens Dal Forno &
Eliasaro.-Figure 44F (Common 7355K,
7367J, 74241).
*Phaeographis ln,,mpic. ,i (Fee) Miill. Arg.-
Figure 44G (Lay 09-01 r1!,). This appears to
be the correct name for the species commonly
identified as P. :,!-ii:Jiii. i (Harris 1995). The
latter is now considered a synonym of P.
brasiliensis (Archer 2009).
Phao.graihli intricans (Nyl.) Staiger-Figure
44H (Common 7355A, '.. ,ey & Seavey
17''_ **).
*Phaeographis irii ,::iiiili (Kremp.) Miill.
Arg.-Figure 45A (Common 7277C, 7355F,
7424E). This is the same taxon asHarris 18150B
in Harris (1995). There are two other species
in Florida (not yet found at Fakahatchee) with
small, muriform ascospores and norstictic acid,
but with inspersed hymenium: Ph.,i. ,. ,i-l,.
multicolor R. C. Harris (with orange hymenium)
and Ph'. i. ;. l,i.. ii'.i J ., j.t (Nyl.) Staiger,
which in Harris (1995) keys out as Small 10114.
There also seems to be an undescribed species
from Florida with clear hymenium and smaller
ascospores which requires further study.
Ph, 'r, jresii lobata (Eschw.) Mull. Arg.-Figure
45B (Common 7423 1).
*Phaeographis major (Kremp.) Liicking-Figure


45C (Common 7355B, 7367J).
*Phaeographis ,rlan,:.ri (Vain.) Zahlbr.-
Figure 45D (Common 7346A, Lay 09-; ")
*Phaeographis scalpturata (Ach.) Staiger-
Figure 45E (Common 7264A, Seavey & Seavey
10297).
Phr 'i::r, jhfl, aff. scalpturata (Ach.) Staiger-
Figure 45F (Common 7381B, Liicking & RA .,,
FL a, 1267 il', 26741). We have not found a name
for this taxon which resembles P!i. "';.''.;i /
scalpturata but has a completely carbonized
excipulum.
*Phrf,tr, .g~iiiphl schizoloma (Miill. Arg.) Miill.
Arg.-Figure 45G (Common 7355C, 7367J,
7381C, Lay 09-0146, i;, ki;- & Rivas Plata
26742).
rPh~.:n':rjfthi. aff. schizoloma (I~Iil. Arg.) Miill.
Arg.-Figure 45H (Common 7262, Lay 09-
0175, Liicking & Rivas Plata 267-1 /', 26746).
This material is the same as Wilson 1259 in
Harris (1995). It agrees with ph4,,,.-i ..,i,..
schizoloma in all details except the lack of
lichen substances and larger ascospores. No
name was found for this taxon, but revision
of types of Ph. ,,',,,- -ii.. has not yet been
concluded.
Phi 'i.:rjfhi, suhfihlgi;mrim (Nyl.) Zahlbr.-
(Buck 54432).
Phra,, .:rijfhi. spec.--(.',', ,hib-' 7624, Hodges
s.n.). We have not found a name for this species.
It is intermediate between P brasiliensis,
in having much branched lirellae with non-
carbonized base, 3-septate ascospores, and
norstictic acid, and .',,. ,. .'.1.. tricosa, in
having an inspersed hymenium.
Phflllfynr, confusa Swinscow & Krog-Figure
46A (Lay 09-0209, 09-0210, Nelsen ii','.)
Plfyt',~,n,.r fitJfitrJ..'a Zahlbr.-Figure 46B
(Lendemer 15699).
Pflt'llT r.," iilinyl, (Vain.) Riddle-Figure 46C
(Crane ILLS *., /37, Lay 09-0106, Liicking &
Rivas Plata 267i '', Seavey & Seavey lo; / ).
Phyllopsora iAailri Brako-Figure 46D (Liicking
& Rivas Plata 26750*).
*Phyllopsora lacerata Timdal-Figure 46E
(Sanders s.n.). This species was recently






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. r',I 1


described from Peru and is also known from
Cuba, Tobago, and Eduador (Timdal 2008),
so its presence in subtropical Florida is not a
surprise. It somewhat resembles E-...hi., i:n..i
but differs in thallus anatomy.
Phyllopsora santensis (Tuck.) Swinscow &
Krog.-Figure 46F (Lay 09-0211).
Physcia atrostriata Moberg-Figure 46G
(Beeching 7711, Buck 54441, Lendemer 15661,
L, .A,/- & Rivas Plata 26751*, 26752, 26753*,
Nelsen i-~,,(., Seavey & ',. ey 1I ',/).
Physcia undulata Moberg-Figure 46H
(Lendemer 15655).
Platygramme .li'i'rnitc ,n (Fee) Fee-Figure
47A (Beeching 7627, Common 7313G; Lay 09-
0107, Lendemer 15693, Lficking & R;, ,. I' h'il..i
26624*, 26826b). The species of Platygramme
are treated in Staiger (2002) and Tripp &
Lendemer (2010).
Platygramme aff. a,, ni'Iriimaii (Fee) Fee-
Figure 47B (Common 7259E, 7292A, 7367K,
7410F, Lay 09-014 '). This material differs from
P caesiopruinosa in the brown, thinly pruinose
disc (bluish pruinose in P .. .. i. i.. i ;) and
the usually pale to almost white thallus.
Platygramme pachnodes (Fee) E. Tripp &
Lendemer-Figure 47C (Seavey & Seavey
10300*). Not in checklist but reported by Tripp
& Lendemer (2010).
Platygramme praestans (Miill. Arg.) Staiger-
Figure 47D (Lucking & Rivas / 'L.., 26628*,
Seavey & Seavey 10586). Not in checklist but
reported by Tripp & Lendemer (2010).
Prla thd,~ iiu flr l.rhi mnt (Tuck.) Lendemer-
Figure 47E (Common 7368P).
Platj ih, iiHm grammitis (F6e) Staiger-Figure
47F (Lendemer 15626).
Polymeridium albocinereum (Kremp.) R. C.
Harris-Figure 47G (Lay 09-0143).
Polymeridium cataphitiin (Nyl.) R. C. Harris-
(Harris & Buck s.n.).
Polymeridum proponents (Nyl.) R. C. Harris-
Figure 47H (Seavey & Seavey 10302*).
Porina heterospora (Fink ex J. Hedrick) R. C.
Harris-Figure 48A (Buck 54414, Lendemer
15562). This species is similar to P nucula,


but differs in the larger ascospores with more
numerous septa (9-13) which are distinctly
tapering towards the proximal end; ascospores
in P nucula are consistently 7-septate and
symmetrically fusiform (Harris 1995).
Porina nucula Ach.-Figure 48B-F (Beeching
7689, Lay 09-0155, Licking & Rivas Plata
26761*, 26762*, Mercado-Diaz 4- 1,, Nelsen
4157, Seavey & Seavey 1Ps" ;' ).
Pseudoparmelia uleana (Miill. Arg.) Elix &
T. H. Nash-(B,--. -hJii 77r0 Buck 54 :.-,
He i, u; i.,*a/ 10552, Lendemer 15563, Seavey &
Seavey 1i 70 /*).
*Pseudopyrenula subgregaria Miill. Arg.-Figure
48G (Licking & Rivas Plata 26770, Nelsen
-4li 72, Seavey & Seavey 10153). Reinstated
for the North American checklist. This taxon
differs from P. diluta (Fee) Mill. Arg. in the
smaller ascospores (20-25 pm versus 25-32
ipm) and the .11, , inspersed hymenium (clear
in P diluta). The thallus is also more compact
and the perithecia flatter than in P diluta.
*Pseudopyrenula subnudata Miill. Arg.-Figure
48H (Lendemer 15656). Reinstated for the
North American i-i- r This taxon differs
from P diluta in the smaller ascospores and the
largely endoperidermal thallus.
Pseudosagedia cestrensis (E. Michener) R. C.
Harris-(Lay 09-0187).
Pseudosagedia aff. cestrensis (E. Michener) R. C.
Harris-(Lay 09-0225, Liicking & Rivas Plata
26757). This material has longer ascospores
with more numerous septa than typical P
cestrensis ascosporess regularly 7-septate).
"Pl, r:ll" spec.-Figure 49A-B (Lay 09-0033,
09-0218, L i..ml;r & Rivas Plata 2I',' n ). In
spite of careful search we have not yet found
a name for this diminutive but characteristic
taxon. The thallus consists of very small (0.1-
0.3 mm diam.), discrete, adnate squamules and
bears abundant, orange-brown to red-brown
apothecia with darker, persistent margin. As-
cospores are non-septate or 1-septate and
12-20 x 1.5-2 pm large, resembling those of
I'i!i 1i, ,i.,. Eventually the squamules break
up to produce large, pale yellow-green soralia.






LUCKING ET AL.: Lichens of Fakahatchee t1.i i Preserve State Park


The taxon comes closest to Psorella pertexta
(Nyl.) Mull. Arg. in the key given by Ekman
(1996: 56), but that species has much longer
ascospores and a F'l li ',I., 'i.-like prothallus
and lacks soralia (Swinscow & Krog 1981).
Another superficially similar species is the
sorediate, microsquamulose Biatora fallax
Hepp (Printzen 1995), which differs in having
Biatora-type apothecia with evanescent margin
and much broader, ellipsoid ascospores. The
fpri-; is here tentatively filed under Psorella
since the type of that genus appears to belong
in Bacidia (S. Ekman, pers. comm. 2011).
*Psoroglaena costaricensis Henssen-Figure
49C-D (Common 7363D).
Punctelia rudecta (Ach.) Krog.-(Beeching
7652).
Pyrenula acutalis R. C. Harris-Figure 49E
(Breuss 28797*, Seavey & Seavey 13333*).
The genus Pyrenda is keyed out for Florida in
Harris (1995).
Pyrenula anomala (Ach.) Vain.-Figure 49F
(Beeching 7654, Lay 09-0153, Liicking &
Rivas Plata s.n., Mercado-Diaz 431, Nelsen
," -3, Seavey & Seavey 10317*).
Pyrenula aspistea (Ach.) Ach.-Figure 49G
(Breuss 28679*, Lay 09-015 1).
Pyrenula astroidea (F6e) R. C. Harris-Figure
49H (LiLcking & Rivas Plata 26774*, Seavey
& Seavey 10322).
'Pj r-iijiii brunnea Fee-Figure 50A (Licking &
Rivas r'li;.. 26777, Seavey & Seavey 10391).
Pyrenula concatervans (Nyl.) R. C. Harris-
Figure 50B (B.. ...hin 7686, Breuss 28993*,
Buck 54427, Lay 09-0157, Licking & Rivas
Plata 26778, Seavey & Seavey 13334*).
Pyrenula confinis (Nyl.) R. C. Harris-Synonym
Pyrenula corlicata (\ lull Arg.) R. C. Harris-
Figure 50C (Common 7 .. IG, 7326M).
Pyrenula cruenta (Mont.) Vain.-Figure 50D
(Beeching 7649, Lay 09-0158, Lendemer
15519, L,.. l: & Rivas Plata 26781, Seavey
& Seavey 10151*).
Pyrenula cubana (Mill. Arg.) R. C. Harris-
Figure 50E (Beeching 7664, Lay 09-0160,
L,,i.,/:. & Rivas P'a,.i. 26783*, 26784*,


Seavey & ',,. i,ey 1I, : .,).
*Pyrenula dermatodes (Borrer) Schaer.-
(Licking & Rivas Plata 26509c). This species
is characterized by a corticate, UV+ yellow
thallus (lichexanthone), immersed perithecia
covered mostly by a thalline layer except for
the ostiolar area, non-inspersed hymenium,
and ascospores 14-20 x 5-6 pm with well-
developed endospore and the terminal lumina
separated from the exospore by an endospore
layer. The species is regularly found in tropical
montane situations and is also known from
Europe; it is therefore surprising that it had not
been reported before from North America.
Pyrenula duplicans (Nyl.) Aptroot-Figure 50F
(In ~, :; & Rivas Plata 267 ,2).
Pyrenula fl.da hin (Zahlbr.) R. C. Harris-Figure
50G (Lay 09-0161, L;,,, n- & Rivas Plata
26786).
Pyrenula globifera (Eschw.) Aptroot-Figure
50H (Breuss 28709*).
Pyrenula leucostoma Ach.-Figure 51A (Crane
ILLS 60469, Lay 09-0162, Lendemer 15688,
Licking & Rivas Plata 26789, 26790*, 26791,
Nelsen i0-7, Seavey & Seavey 10307*).
Pyrenula mamillana (Ach.) Trevis.-Figure 51B
(Lay 09-0163, Liicking & Rivas Plata 26792,
26795*).
Pyrenula microtheca R. C. Harris-Figure 51C
(Common 7326B, Lay 09-0109, Seavey &
Seavey 10 i)
Pyrenula mucosa (Vain.) R. C. Harris-Figure
51D (Licking & Rivas Plata 26797*, 26798*,
26812, Nelsen 4096).
Pyrenula hi,, -rfli ii (Nyl.) R. C. Harris-
Figure 51E (~...,.air 7590, Crane ILLS
'' i79, Mercado-Diaz -, .1, Nelsen ,i.,*.,
Seavey & Seavey 10309).
Pyrenula ochraceoflavens (Nyl.) R. C. Harris-
Figure 51F (Seavey & Seavey 10311).
Pyrenula punctella (Nyl.) Trevis.-Figure 51G
(7 n./, l,: & Rivas Plata 26801).
Pyrenula qiii;,'i, a l F6e-Figure 51H (Beeching
7599, Lay 09-0166, Lendemer 15689, L,_/.r'-
& Rivas Plata 26803, Mercado-Diaz 432,
Seavey & Seavey 10519).






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P,' 1,


Pyrenula santensis (Nyl.) Miill. Arg.-Figure 52A
(Lay 09-0170, Licking & Rivas Plata 26785,
2,,'i i, 26805, Seavey & Seavey 10536).
P ru.,.-ilai aff. santensis (Nyl.) Miill. Arg.-Figure
52B /; ', /ni,-Diaz 442). This material is the
same as the one cited in Harris (1995: 106) for
Collier County under the collection number
Harris 30313. It agrees with P. santensis in
the large perithecia covered by a thin thalline
layer and the non-inspersed hymenium, but has
smaller ascospores (14-15 x 4.5-5.5 pm in the
present material). We agree with Harris (1995)
that this might represent a taxon distinct from
P santensis s.str.
Pyrenula .,?qi, llm'ii (F;ihw.) R. C. Harris-
(_;../ ,, ,,./i. Seavey & Seavey 13335*).
*Pyrenula sexlocularis (Nyl.) Miill. Arg.-Figure
52C (Liicking & Rivas Plata 26780, Seavey
& Seavey l"' ,* ). This taxon has almost
invariably been included in P concatervans,
a species with 3-septate ascospores (Harris
1995). Considering that ascospores with more
than three transverse septa are extremely rare
within Pyrenula, and the material seen by
us has either 3-septate or 5-septate mature
ascospores, we propose to recognize the taxon
with 5-septate ascospores as separate species.
P6 r,..evil thelomorpha Tuck.-Figure 52D
(Liicking & Rivas PF, .' 26658b, Seavey &
Seavey l',', i).
Pyrgillus javanicus (Mont. & Bosch) Nyl.-
Figure 52E (f.v,, ,:;;hi 7,.'i i, Lay 09-0110,
Lendemer 15583).
Pyxine cocoes (Sw.) Nyl.-Figure 52F (Licking &
R' .Plata 26806).
Pyxine eschweileri (Tuck.) Vain.-Figure 52G-
H i,' *,'... 28655*, Lay 09-0060, Liicking &
Rivas Plata 26563b, Mercado-Diaz ,i / ).
Ramalina complanata (Sw.) Ach.-(H .i,,i ,
10547, Seavey & Seavey lo i26).
Ramalina dasypoga Tuck.-(Lay 09-0111, Seavey
& Seavey 10579*).
Raimnlina dendriscoides Nyl.-(Lendemer 15690).
Ramalina denticulata Nyl.-(Lay 09-0112, Seavey
& Seavey 10427).
Ramalina nreleIre,:.i De Not.-(E.,-'.. /,hi;, 7610,


Breuss ',' /*, McMullin 301 '..
Rantlina peruviana Ach.--(Z P,'.-', i;,, 7634,
Seavey & Seavey 1041- ).
Rainutlina .,mr t .. Miill. Arg.-(Crane ILLS
,-- t 1-, Lay 09-0113, Seavey & Seavey 10835).
Randlina usnea (L.) R. Howe-(Beeching 7616,
Crane ILLS '-/ 38, H ,',i,..ln 10556, Lay
09-0116, Lendemer 15593, Seavey & Seavey
I, ',. /*).
Ranuilina willjy R. Howe.-(f., 'i.. hiw 7719).
Reimnitzia santensis (Tuck.) Kalb-Figure 53A-
B (Liicking & Rivas Plata 26558*, Nelsen
4179).
Sarcographa bth.driti_.,n (Ach.) Miill. Arg.-
Figure 53C-H (Beeching 7635, Buck 54461,
Common 7355G, 7367A, 7381A, Crane ILLS
"1,,t i0, Lay 09-0119, Lendemer 15678, Liicking
& Rivas Plata 26559*, 26560*, Seavey &
Seavey 10799*). .' ,. [. 1 / ;,.7, i; i, i 1 is
possibly a collective species. The shape and
arrangement of the lirellae within the stromata
is very different across samples (Figure 53C-
H) and this appears to correlate with thallus
morphology.
Sri,,i.gr, q, pa tricosa (Ach.) liill. Arg.-Figure
54A (Common 7346H, 7367F, 7410D, Crane
ILLS o; I Hodges 9304.7, Lay 09-0147).
Segestria lctgldc.. (Durieu & Mont.) R.C.
Harris-Figure 54B () ,', ,.. 28722*).
*Sporopodium marginatum Liicking &
Lumbsch-Figure 54C (Liicking & Rivas
Plata 2't,.:.,'.., ;:, /,/,- 86, 87, 90, 91).
i,:;, ,,hr\ auberianus (Mont.) A. Frisch &
Kalb--(EB.','. 1hi<; 7690).
i,~:d,,iru\ emersus (Kremp.) A. Frisch &
Kalb-Figure 54D (Harris & Buck s.n). In the
checklist as O, l7 J'. ,/ i. emersa.
Stegobolus ,,nuir lum\ (Tuck.) A. Frisch.-
Figure 54E-H (Lay 09-0028, Lendemer 15602,
Mercado-Diaz .iou, Nelsen 4180, Seavey &
Seavey 1r. :: 7).
Sticta beauvoisii Delise-(Beeching 7655, Lay 09-
0121).
**\irmimini dubia A. L. Sm.-Figure 55A-B
(Common 7421D). The genus Stirtonia was
revised by Aptroot (2009), who stated that the






LUCKING ET AL.: Lichens of Fakahatchee t.I i Preserve State Park


genus is almost absent from the Neotropics.
Surprisingly, two species previously known
only from the Paleotropics were found in
our material, indicating that this genus is
probably undercollected in the Neotropics or
misidentified in herbarium collections.
**Stirtonia macrocarpa Makhija & Patw.-
Figure 55C-D (Common 7327C).
*1:iil:,ii orbicularis Fr.-Figure 55E (Seavey &
Seavey 10510).
.Vii7j iiLi phaea (Ach.) R. C. Harris-Figure 55F
(Buck 54474, Lendemer 15531, Lu .Aur- &
Rivas Plata 2'. -' )).
*Strigula schizospora R. Sant.-Figure 55G (Lay
09-0208A, Mercado-Diaz 419a, ',. tiranek 109).
Strigula iriau.',hfhLr Fr.-Figure 55H (Lay
09-0208B, Licking & Rivas Plata 26694b,
Mercado-Diaz 419b, Nelsen 4077, Seavey &
',. ,ey 10531).
S}l cd.,i' byssina (Vain.) Tehler-Figure 56A
(Breuss 28938*, Lendemer 15619, Liicking &
Rivas J'ai.. 26831*, Seavey & Seavey 10781).
*Tapellaria albonarginata Liicking-Figure 56B
(Common 7_'....F, 7322H, Seavey & Seavey
10571). This species is described in a recent
paper (Lumbsch et al. 2011) based on material
from Costa Rica. It is anatomically similar
to T bilimbioides R. Sant. but differs in the
distinct white pruina covering the apothecial
margin. The ascospores are predominantly
3-septate but occasionally, ascospores with up
to four or five septa might occur. In the North
American checklist, this name should replace
T. .T'!,id ,. which is with certainty only
known from the Paleotropics.
****T;Yj', llin- jlJidk'f, iCommon & Liicking-
Figure 56C (Common 7315D, Common
7322A). See p. 149 for description of this new

***T""jY ll .I granulosa Liicking & Rivas
Plata-Figure 56D (1., 09-0013, Liicking
& Rivas LJ.ia.. 26697, 26810*, Mercado-Diaz
;' !). See p. 149 for description of this new
species.
*Tapellaria malmei R. Sant.-Figure 56E (Lay
09-0017B, LI.. r, ; & Rivas Plata 26811).


T, filawia nana R. Sant.-Figure 56F (Seavey &
Seavey 10618).
Tpriwl,wir a spec.-Figure 56G-H (Hodges s.n.).
This material is characterized by a coarsely
verrucose-bullate thallus; the hypothecium and
excipulum are dark gray-brown reacting K+
sordid green, a reaction unusual for the genus
(normally K+ purple). While campylidia are
abundant in the material, only one apothecium
was found and it lacked ascospores. Thus,
while this taxon is certainly not identical with
any known species of T.q,,-, ..'i..l it cannot be
formally described at this point, but further
collections must be awaited to get ascospore
data.
Tphj'lr,,r,:tr atra (Hudson) Hafellner-Figure
57A (Seavey & Seavey 10574*).
Thalloloma me.;iiirmI;ri (Mont.) Trevis.-Figure
57B (Common 7356K, 7367B, 7.:'- /).
Thalloloma hypoleptum (Nyl.) Staiger-Figure
57C (Liucking & Rivas Plata 265':- / ).
Thecaria quassiicola F6e-Figure 57D (B,-,: ,/iS
7661, Buck 54477, Common 7313B, 7366,
Crane ILLS a i 1, Lay 09-0117, Licking &
Rivas Plata 26615*, 26626*, Seavey & Seavey
10603). The species is extremely variable in
producing both perfectly round and distinctly
lirellate ascomata on the same thallus (Harris
1995). Exactly the same variation was
observed in material from the Philippines and
the -.p'" iiC, i... cluster together in molecular
phylogenetic analyses (Rivas Plata et al.,
unpubl. data).
Thelotrema lathraeum Tuck.-Figure 57E
(Common 7359B, 7371A, Harris & Buck
s.n.). This species was synonymized with T.
.lj;,, ,.::; (Mull. Arg.) Mangold (Rivas Plata
et al. 2010), but is actually set apart by the
distinctly corticate thallus and extremely small
apothecia. It frequently grows together with
Fissurina ,i,,., 1.,',.,'//,., which has the same
thallus type.
*Thelotrema /ipa~ ,h \,r'i m Nyl.-Figure 57F
(E.,, i-h,1t- 7628, I n, ,: & Rivas Plata
26569*, Seavey & Seavey 10812).
Thelotremai p'rimi;dl.'. Mont. & Bosch.-Figure






BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. P', 1,


57G (7 il. i Iw,- & Rivas Plata 26570).
Thelotrema ,ihiidL' Tuck.-(Beeching 7723).
Tricharia subumbrosa Liicking & W. R. Buck.-
i ./:,',. ;1111 7700).
Tricharia vainioi R. Sant.-Figure 57H (Crane
ILLS 60435, Lnl.,,: & Rivas Plata 2<...'.,,
Mercado-Diaz 419c).
T.j,'iheliFum aeneum (Eschw.) Zahlbr.-Figure
58A iT....;.,- 7639, 7696, Buck 54430,
Crane ILLS 60459, Lendemer 15695, Lciiking
& Rivas Plata 26814*, Nelsen e,'1, Seavey &
Seavey 10514*).
T ejctheilium eluteriae Spreng.-Figure 58B
i7., .../,,:, 7680, Mercado-Diaz 434, 452,
Seavey & Seavey 10532*).
Trij'thliiufit mnarcidum (Fee) Aptroot-Figure
58C Synonym T ji i 1,.,,,,,,, (Zahlbr. ex
Choisy) R. C. Harris-(B!.,, hi 7694, Buck
54393, Lendemer 15543, Mercado-Diaz
479, Nelsen w'o, Seavey & Seavey 10587).
As Ti j'.. i,.ii/ ,m floridanum in the North
American checklist.
T',yJ''thlinu h nitidiusculum (Nyl.) R. C.
Harris-Figure 58D (', .ii, hi;.* 7692, Crane
ILLS 60466, Lay 09-0122, Liicking & Rivas
Plata 26816*, Nelsen 4-i_'a, Seavey & Seavey
1,' 118).
TJ'&,y'th,'li,& ochroleucum (Eschw.) Nyl.-
Figure 58E (P:,', !;i,.: 7709, Crane ILLS
60452, Lay 09-0144, Licking & Rivas Plata
26822*, 26824*, Mercado-Diaz 417, Nelsen
4081, Seavey & Seavey 10161*).
TJ',t'th,'linun tropicum (Ach.) Miill. Arg.-
Figure 58F (P., .!'io: 7695, Buck 5 / ', Lay
09-0125, Lendemer 15581, Liicking & Rivas
Plata 26827, 2,;.'8*, Nelsen 4143, Seavey &
Seavey 10 li1*).
Tylophoron moderatum Nyl.-Figure 58G-H
(Harris & Buck s.n.).

ACKNOWLEDGMENTS

We thank the Fakahatchee Strand Preserve State
Park staff and volunteers, in particular Mike
Owen, Donna Glann-Smyth, Bob Nesmith, Alicia
Campanella, Karen Relish, Laura Skinner, and


Richard Fagan, for their support of this project and
especially for their i-'j.-wim- and knowledgeable
guidance through the park during our stay. The
Florida Department of Environmental Protection,
Division of Recreation and Parks, is kindly
acknowledged for issuing collection permits in
1997 to Ralph Common 1-:1 1169714), in 2009 to
William Safranek for the Tuckerman workshop
(#4-09-02), and in 2010 to William Sanders (#4-
10-34). Thanks also to Martha J. Robinson, Florida
Department of Environmental Protection, for the
p:i ,iii-,.,-,, to use the park map depicted in Figure
2. Linda Weinland organized the perfect laboratory
facilities at Edison State College. Klaus Kalb
and Einar Timdal assisted with the identification
of C i. db! i '/, J ,un. endochroma and P/in 7if.' ,
lacerata, and Andre Aptroot gave valuable advice
regarding some of the pyrenocarpous taxa. The
Tuckerman workshop was partially funded by the
NSF project "Neotropical Epiphytic Microlichens
- An Innovative Inventory of a Highly Diverse
yet Little Known Group of Symbiotic Organisms"
(NSF-DEB 0715660 to The Field Museum; PI R.
Lucking). Part of the identifications were done
within the framework ofphylogenetic studies of the
lichen family Graphidaceae, supported by two NSF
grants: "Phylogeny and Taxonomy of Ostropalean
in-," (DEB 0516116; PI Lumbsch, Co-PI
Lacking) and "ATM Assembling a taxonomic
monograph: The lichen family Graphidaceae"
(DEB 1025861; PI Lumbsch, Co-PI I iki iA,).
Curators of the herbaria in B, BM, CANB, FH,
G, H, M, NY, S, TUR, UPS, US, and W provided
loans of, or access to, valuable type material and
other significant collections.

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Tripp, E. A., J. C. Lendemer, & R. C. Harris.
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Muill. Arg. in North America: new species,
new combinations, and treatments for
.-4,, ,;ubi .. ('o -.-.... i .1, / .and
Di, ,;,;. .'. Lichenologist 42:55-71.
Tucker, S. C. 1979. New or noteworthy records
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82:125-140.
Tucker, S. C. 1981. Checklist of Louisiana lichens.
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Tucker, S. C. 2010. Lichens of Burden Research
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Wetmore, C. M. 1994. The lichen genus C./, ,/*.. ,l
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Contributions to Botany 40:1-64.






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Figure 1. Group photograph of the 2009 Tuckerman workshop. Standing from left to right: Lee
Crane, Sean Beeching, Frederick Seavey, Malcolm Hodges, Richard Fagan, William Buck,
Elisabeth Lay, Mike Owen, Harold Schaefer, Matthew Nelsen, William Sanders, Troy
McMullin, and Brendan Hodkinson. Kneeling from left to right: Jean Seavey, Bob Nesmith,
Alicia Campanella, James Lendemer, William Safranek, Ottmar Breuss, Joel Mercado-Diaz,
Eimy Rivas Plata, Natasha Lucking Rivas Plata, and Robert Lucking.











Fakahatchee
Strand Preserve.
State Park J__ '

Collier County.

A 2 mi

Figure 2. A, County map of Florida showing Collier County and the study site. B, Map of
Fakahatchee Strand Preserve State Park, straddling the Tamiami Trail (Route 41) within Collier
County; the four main collection sites described in the text are indicated by numerals 1 to 4. Map
reprinted with permission of the Florida Department of Environment Protection, Tallahassee,
Florida.

From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.





















































Figure 3. Habitat photographs of the four collection sites at Fakahatchee Strand Preserve State
Park. First row: site 1, showing royal palm canopy and understory with bald cypress. Second
row: site 2, showing side road along old tram with dense vegetation and aspect of understory
with bromeliads. Third row: site 3, showing open bald cypress prairie hammock and dense
regrowth. Fourth row: site 4, showing closed bald cypress forest and aspect of forest understory.
Photographs by R. Lacking (A, B, G), Matthew Nelsen (D, E, H), Frederick Seavey (F), and Jean
Seavey (C).








From on-line supplemental materials to:
Licking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.










SUPPLEMENTAL MATERIALS

FIGURES 4-58

Liicking, R., F. Seavey, R. S. Common, S. Q. Beeching, O. Breuss, W. R. Buck, L. Crane, M.
Hodges, B. P. Hodkinson, E. Lay, J. C. Lendemer, R. T. McMullin, J. A. Mercado-Diaz,
M. P. Nelsen, E. Rivas Plata, W. Safranek, W. B. Sanders, H. P. Schaefer Jr. & J. Seavey.
2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from
the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History
49(4):127-186.

Copyright for individual images belongs to the listed photographer.









































*' -- l '

; ... e.. :..
". .* . H.g-
-, .- <,o ,w
.4 1.Y
: c;."~L4


Figure 4. A-B, Acanthothecis peplophora (A, Lfcking 26501; B, Seavey & Seavey 11007). C, Acanthothecis
poitaeoides (Seavey & Seavey 10843). D, Actinoplaca spec. (Sanders 10504.6). E, Amandinea endachroa (Breuss
28936). F, Amandinea punctata (Seavey & Seavey 10712). G, Anisomeridium subnexum (Lfcking & Rivas Plata
26616). H, Anisomeridium subprostans (Seavey & Seavey 10700). Photographs R. Liicking (A-B, D-E, G), R.
Seavey (C, F, H).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.
































H-of
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Figure 5. A, Anisomeridium spec. (L"cking & Rivas Plata 26598). B-D, Anthracothecium prasinum (Lcking &
Rivas Plata 26595, 26594, 26292). E-F, Arthonia antillarum (Common 7327E). G-H, Arthonia cinnabarina
(Common 7421C). Photographs R. Common (E-H), R. Lcking (A-D).
From on-line supplemental materials to:
LfRcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4): 127-186.


5 ir
























































Figure 6. A, Arthonia complanata (Seavey & Seavey 10542). B. Arthonia ilicina (Mercado-Diaz 410). C-D,
Arthonia interveniens (Common 7382A). E-F, Arthonia macrotheca (Seavey & Seavey 10167). G-H, Arthonia
mirabilis (Common 7265). Photographs R. Common (C-D, G-H), R. Liicking (B), R. Seavey (A, E-F).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.
~~-4-





3-4














Tuckennan Workshop. Bulletin of the Florida Museum of Natural History 49(4): 127 186.































































Figure 7. A, Arthonia aff. pinastri (Lay 09-0042). B, Arthonia platygraphidea (Licking & Rivas Plata 26583). C-
D, Arthonia aff. pruinata (Common 7372A). E-F, Arthonia pruinosella (Common 7327A). G-H, Arthonia
pyrrhuliza (Lay 09-0017A). Photographs R. Common (C-F), R. Liicking (A-B, G-H).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.










IF








-f J


I. -


Figure 8. A-B, Arthonia redingeri (Common 7259U). C, Arthonia rubella (Licking & Rivas Plata 26586). D,
Arthonia rubrocincta (Lfcking & Rivas Plata 26582). E-F, Arthonia simplicascens (Common 7291D). G-H,
Arthonia speciosa (Common 72911). Photographs R. Common (A-B, E-H), R. Liicking (C-D).

From on-line supplemental materials to:

Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.
















































i:M


Figure 9. A, Arthopyrenia cinchonae. B, Arthopyrenia lyrata. C, Arthopyrenia planorbis (Seavey & Seavey 10165).
D, Arthopyrenia aff. planorbis (Licking & Rivas Plata 26787b). E, Arthothelium spectabile (Breuss 28770). F,
Aspidothelium cinerascens. G, Aspidothelium geminiparum (Lfcking & Rivas Plata 26600c). H, Aspidothelium
scutellicarpum (Breuss 28787). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.































































Figure 10. A, Astrothelium confusum (Licking & Rivas Plata 26691). B, Astrothelium diplocarpoides (Licking &
Rivas Plata 26627). C, Astrothelium diplocarpum (Nelsen). D, Astrothelium galbineum (Lfcking & Rivas Plata
26596). E-F, Astrothelium variolosum (Breuss 28962). G, Aulaxina microphana (Sanders 10521.1). H, Aulaxina
quadrangula, hyphophores (Sanders 10521.7). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.











































































Figure 11. A, Bacidia aggregatula (Seavey & Seavey 10095). B, Bacidia campalea (Breuss 28735). C, Bacidia
heterochoa (Licking & Rivas Plata 26621). D, Bacidia aff. heterochroa (Licking & Rivas Plata 26610). E. Bacidia
hostheleoides (Licking & Rivas Plata 26605). F, Bacidia aff. hostheleoides (Licking & Rivas Plata 26600b). G,
Bacidia medialis (Licking & Rivas Plata 26601). H, Bacidia aff. medialis (Licking & Rivas Plata 26606).
Photographs R. Lacking (B-H), R. Seavey (A).

From on-line supplemental materials to:

Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


*


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C~r~i:ir
I rr.le
Ta: :'S;
rlri

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Figure 12. A, Bacidia mutabilis (Licking & Rivas Plata 26603). B, Bacidia russeola (Breuss 28828). C, Bacidia
schweinitzii (Seavey & Seavey 13331). D, Bacidia aff. schweinitzii (Licking & Rivas Plata 26602a). E, Bacidia
spec. (Licking & Rivas Plata 26611). F, Bacidina varia (Lay 09-0004). G, Bactrospora denticulata (Licking &
Rivas Plata 2673 7b). H, Bactrospora myriadea (Lay 09-0212). All photographs R. Liicking.

From on-line supplemental materials to:

Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


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''


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.~.
































































Figure 13. A, Baculifera curtisii (Seavey & Seavey 10432). B, Baculifera imshaugiana (Breuss 28890). C,
Bathelium carolinianum (Nelsen 4149). D, Bathelium madreporiforme (Lay 09-0047). E, F, o;lh,,-.... leucoxantha
(Breuss 28981). F, Byssoloma chlorinum (Nelsen). G, Byssoloma leucoblepharum (Lfcking & Rivas Plata 26692).
H, Byssoloma meadii (Lendemer 15572). Photographs R. Liicking (B-H), R. Seavey (A).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 14. A-B, Calicium hyperelloides (Seavey & Seavey). C, Calopadia editae (Common 7322B). D-G,
Calopadia floridana (D-E, holotype; F-G, Common 5320K). H, Calopadia fusca (Seavey & Seavey 10365).
Photographs R. Common (C, F-G), R. Liicking (D-E), R. Seavey (A-B, H).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.








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,*, L'' ., '




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Figure 15. A-F, Calopadia imshaugii (A-B, E, holotype; C-D, Homestead, Common 58891; F, Homestead,
Common 5892E). G-H, Calopadia lecanorella (Nelsen s.n., Licking & Rivas Plata 26696). Photographs R.
Common (A-F), R. Liicking (G-H).

From on-line supplemental materials to:

Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


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Figure 16. A, Calopadia perpallida (Nelsen). B, Calopadia puiggarii. C, Calopadia subcoerulescens. D, Caloplaca
camptidia (Lucking & Rivas Plata 26568b). E, Caloplaca epiphora (Seavey & Seavey 10543). F, Caloplaca
holocarpa (Lfcking & Rivas Plata 26698). G, Catillochroma endochroma (Beeching). H, Celothelium aciculiferum
(Common 7396D). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


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Figure 17. A, Chaenotheca brunneola (Lay 09-0049). B, Chapsa chionostoma (Common 7321A). C, Chapsa
platycarpa (Lficking & Rivas Plata 26573). D, Chapsa platycarpoides (Lficking & Rivas Plata 26571). E,
C hi, 'li ,, xanthina (Nelsen). F, Clathroporina isidiifera (Lficking & Rivas Plata 26768). G, Clathroporina
subpungens (Seavey & Seavey 10241). H, Clathroporina tetracerae (Lfcking & Rivas Plata 26767). All
photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 18. A, Coccocarpia domingensis. B, Coccocarpia erythroxyli (Seavey & Seavey 10382). C-D, Coccocarpia
palmicola (Lfcking & Rivas Plata 26701). E, Coenogonium congense (Lay 09-0134). F, Coenogonium geralense
(Lfcking & Rivas Plata 26702). G, Coenogonium interplexum (Lay 09-0132). H, Coenogonium linkii (Lay 09-
0002). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.

































. -4 A --
~1L24Q*? B


Figure 19. A, Coenogonium luteocitrinum (Lay 09-0131). B, Coenogonium luteum. C, Coenogonium subdentatum
(Lay 09-0130). D, Coenogonium subfallaciosum (Lfcking & Rivas Plata 26703). E, Coniarthonia wilmsiana
(Common 7291G). F, Cratiria lauricassiae (Common 7287E). G, Crocynia gossypina (Seavey & Seavey 10473). H,
Crocynia pyxinoides (Licking & Rivas Plata 26705). Photographs R. Common (E), R. Lacking (all others).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.































































Figure 20. A, Cryptolechia nana (Breuss 28711). B, Cryptothecia ettiu..u C-D, Cryptothecia evergladensis
(Lfcking & Rivas Plata 26670). E-F, Cryptothecia miniata (Lfcking & Rivas Plata 26678). G, Cryptothecia
punctosorediata (Lfcking & Rivas Plata 26680). H, Cryptothecia striata (Lfcking & Rivas Plata 26687). All
photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.































































Figure 21. A, Dictyonema ph/lll/oternmnI (Common 7363D). B, Diitrcunim sericeum f. lph.llophlAdrn (Costa Rica,
Liicking s.n.). C, Di'rryiaLI jurnfhlhairh i (Common 7418D). D, Diori gtm microsporum (Liicking & Rivas Plata
26504). E, Diorygma poitaei (Seavey & Seavey 1054.y). F, Dirinaria aegialita (Seavey & Seavey 10621). G,
Dirinaria confusa (Brazil, Cdceres s.n.). H, Dirinaria leopoldii (Seavey & Seavey 10632). Photographs R. Lacking
(A-E, G-H), R. Seavey (F).
From on-line supplemental materials to:
Licking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49t 4.1:127-186.
















































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k. .. -
2'*-* *9 -" *.


Figure 22. A-B, Dirinaria papillulifera (Colombia, Lfcking 32504). C, Dirinaria picta (Seavey & Seavey 10633).
D, Dirinaria purpurascens (Seavey & Seavey 10636). E-F, Dyplolabia afzelii (Lfcking & Rivas Plata 26509,
26505). G, Echinoplaca areolata (Common 7285B). H, Echinoplaca aff. leucotrichoides (Crane). Photographs R.
Licking (A-B, D-H), R. Seavey (C).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 23. A, Echinoplaca lucernifera (Sanders 10504.1b). B, Echinoplaca similis (Breuss 28922). C,
Enterographa anguinella (Seavey & Seavey 10429). D, Eugeniella leucocheila. E, Fissurina aggregatula
holotypee). F-G, Fissurina analphabetica holotypee). H, Fissurina cingalina (Common 7323F). Photographs R.
Common (G), R. Liicking (all others).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.





















L -004


Figure 24. A, Fissurina columbina (Lendemer). B, Fissurina confusa (Common 7356A). C-D, Fissurina
crassilabra (Common 7356C). E, Fissurina egena (Common 7276B, 7368Q). F, Fissurina aff. elaiocarpa (Common
73567). G-H, Fissurina humilis (Homestead, Common 5887F). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


"' '' LI~o;
L~1
*I I~
;L' .. ~L~I.C
* i+
i!
~p~ a :, ir,
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~Fi~i~i~-































































Figure 25. A, Fissurina illiterate (Licking & Rivas Plata 26618). B, Fissurina inspersa holotypee). C, Fissurina
mexicana (Licking & Rivas Plata 26513). D, Fissurina pseudostromatica holotypee). E, Fissurina radiata
(Common 7356A). F, Fissurina rufula (Breuss 28769). G, Fissurina subcomparimuralis holotypee). H, Fissurina
subnitidula (Common 74181). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.































"I'll 'a


Figure 26. A, Fissurina tachygrapha (Common 7418A). B, Fissurina tuckermaniana holotypee). C, Fissurina
varieseptata holotypee). D, Flakea papillata (Lay 09-0062). E, Glyphis atrofusca (Lay 09-0015). F-G, Glyphis
cicatricosa (Common 7259C). H, Glyphis scyphulifera (Common 7292D). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.































































Figure 27. A, Graphis cf. acharii (Licking & Rivas Plata 26521). B, Graphis anfractuosa (Lay 09-0070). C,
Graphis aperiens (Common 7425D). D-E, Graphis appendiculata (D, holotype; E, Licking & Rivas Plata 26657).
F, Graphis argentata (Breuss 28825). G, Graphis assimilis (Everglades City, Common 73461). H, Graphis caesiella
(Everglades City, Common 7341A). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 28. A, Graphis caesiocarpa (Breuss 28718). B, Graphis caribica (Common 7356G). C, Graphis conferta
(Lay 09-0016). D, Graphis cupei (Lfcking & Rivas Plata 26536a). E, Graphis desquamescens (Common 73680).
F-G, Graphis disserpens (Licking & Rivas Plata 26656). H, Graphis handelii (Common 7292G). All photographs
R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


W,,ff ^"i.ow- ".-II.l



















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Figure 29. A, Graphis longula (Lucking & Rivas Plata 26527). B, Graphis lucifica (Lay 09-0065). C, Graphis
oshioi (Lucking & Rivas Plata 26612). D, Graphis oxyclada (Hodges s.n.). E-G, Graphis pseudocinerea Lfcking
(E, Breuss 28987; F-G, Lacking & Rivas Plata 26537). H, Graphis rimulosa (Costa Rica: Standley 71849). All
photographs R. Lfcking.
From on-line supplemental materials to:
Lfcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4): 127-186.































































Figure 30. A, Graphis sauroidea (Breuss 28890). B, Graphis subflexibilis (Common 7380G). C-F, Graphis stellata
(Breuss 28889). G, Graphis xanthospora (Lfcking & Rivas Plata 26535). H, Graphis xylophaga (Seavey & Seavey
10359). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.





























































Figure 31. A-B, Gyalectidium appendiculatum (Sanders 10504.6). C, Gyalectidium catenulatum (Sanders
10504.8). D-E, Gyalectidium floridense (Sanders 10521.4). F, Gyalectidium imperfectum (Sanders 10504.7). G,
Gyalectidium ulloae (Sanders 10504.10). H, Gyalectidium aff. yahriae (Sanders 10521.2). All photographs R.
Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.
















Lh
X t
l^Bi:-


14


Figure 32. A, Haematomma flexuosum (Nelsen). B, Haematomma guyanense. C, Haematomma persoonii (Seavey
& Seavey 10070). D-E, Hafellia bahiana (Everglades: Seavey & Seavey). F, Hafellia curatellae (Seavey & Seavey
10615). G, Hafellia pleiotera (Breuss 28936). H, Halegraphafloridana (Common 7410C). Photographs R. Liicking
(A-B, G-H), R. Seavey (C-F).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.


G I r






























































Figure 33. A-B, Heiomasia seaveyorum. C-D, Herpothallon antillarum (Nelsen 4037). E, Herpothallon echinatum
(Licking & Rivas Plata 26681). F-G, Herpothallon rubrocinctum (F, Licking & Rivas Plata 26707; G, Lcking &
Rivas Plata 26708). H, Herpothallon rubroechinatum (Licking & Rivas Plata 26682). All photographs R. Liicking.
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 34. A, Heterodermia albicans (Breuss 28763). B, Heterodermia pseudospeciosa (Breuss 29020). C-D,
Laurera megasperma (C, L~cking & Rivas Plata 26709; D, Lfcking & Rivas Plata 26710a). E, Lecanora achroa
(Seavey & Seavey 10144). F, Lecanora achroides (Seavey & Seavey 10624). G, Lecanora allophana (Lfcking &
Rivas Plata 26639). H, Lecanora argentata (Licking & Rivas Plata 26634b). Photographs R. Liicking (A-D, G-H),
R. Seavey (E-F).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.






























































Figure 35. A, Lecanora caesiorubella ssp. glaucomodes (Seavey & Seavey 10077). B-C, Lecanorafloridula (Lay
09-0083). D, Lecanora hybocarpa (Lay 09-0138). E, Lecanora leprosa (Breuss 28936). F, Lecanora
pseudargentata (Lficking & Rivas Plata 26642). G, Lecanora strobilina (Lficking & Rivas Plata 26637). H,
Leiorreuma exaltatum (Common 7424A). Photographs R. Lacking (B-H), R. Seavey (A).
From on-line supplemental materials to:
Liicking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18t
Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127-186.




Full Text

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BULLETIN THE LICHENS OF FAKAHATCHEE STRAND PRESERVE STATE PARK, FLORIDA: PROCEEDINGS FROM THE 18 th TUCKERMAN WORKSHOP Robert Lcking, Frederick Seavey, Ralph S. Common, Sean Q. Beeching, Othmar Breuss, William R. Buck, Lee Crane, Malcolm Hodges, Brendan P. Hodkinson, Elisabeth Lay, James C. Lendemer, R. Troy McMullin, Joel A. Mercado-Daz, Matthew P. Nelsen, Eimy Rivas Plata, William Safranek, William B. Sanders, Harold P. Schaefer Jr., and Jean Seavey Vol. 49, No. 4, pp. 127 2011 UNIVERSITY OF FLORIDA GAINESVILLE

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The FLORIDA MUSEUM OF NATURAL HISTORY is Floridas state museum of natural history, dedicated to understanding, preserving, and interpreting biological diversity and cultural heritage. The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is a peer-reviewed journal that publishes results of original research in zoology, botany, paleontology, archaeology, and museum science. The Bulletin is published at irregular intervals, and volumes are not necessarily completed in any one year. Volumes contain between 150 and 300 pages, sometimes more. The number of papers contained in each volume varies, depending upon the number of pages in each paper, but four numbers is the current standard. Multi-author issues of related papers have been published together, and inquiries about putting together such issues are welcomed. Address all inquiries to the Managing Editor of the Bulletin. Richard C. Hulbert Jr., Editor Bulletin Committee Ann S. Cordell Richard C. Hulbert Jr. Jacqueline Miller Larry M. Page Roger W. Portell, Treasurer Irvy R. Quitmyer David W. Steadman, ISSN: 0071-6154 Copyright 2011 by the Florida Museum of Natural History, University of Florida. All rights reserved. Text, commercial use or republication by printed or electronic media is strictly prohibited without written permission of the museum. Publication Date: March 31, 2011 Price: $9.00 This issue completes Volume 49 Send communications concerning puchase or exchange of this publication and manuscript queries to: Managing Editor of the Bulletin Florida Museum of Natural History University of Florida P.O. Box 117800 Gainesville, FL 32611-7800 USA FAX: 352-846-0287

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ABSTRACT Fakahatchee Strand Preserve State Park is located in Collier County at the extreme southwestern corner of Florida, close to Everglades National Park and Big Cypress National Preserve. The 18 th Tuckerman Workshop, an annual gathering of professional and amateur lichenologists and mycologists from the United States and Canada, this time with additional participants from Puerto Rico, Peru, and Austria, was within the Preserve. Together with previously made collections, the survey produced a total of 432 taxa, 18 of which are new to science and 89 are additions to the North America checklist, six of which are also new to the New World. The new species are: Calopadia Hodges & Lcking, Calopadia imshaugii Common & Lcking, Vain. Lcking, M. Cceres & Lcking (formally described in a separate paper), Common & Lcking, Common & Lcking, Common & Lcking, Common & Lcking, Lcking & Rivas Plata, Common & Lcking (formally described in a separate paper), Common & Lcking, Common & Lcking, Common & Lcking Common & Lcking (formally described in a separate paper), M. P. Nelsen & Lcking (formally described in a separate paper), Common & Lcking, Common & Lcking, and Lcking & Rivas Plata. Further, the following three new combinations are proposed: Chapsa (Tuck.) Breuss & THE LICHENS OF FAKAHATCHEE STRAND PRESERVE STATE PARK, FLORIDA: PROCEEDINGS FROM THE 18 th TUCKERMAN WORKSHOP Robert Lcking 1,20 Frederick Seavey 2 Ralph S. Common 3 Sean Q. Beeching 4 Othmar Breuss 5 William R. Buck 6 Lee Crane 7 Malcolm Hodges 8 Brendan P. Hodkinson 9 Elisabeth Lay 10 James C. Lendemer 11 R. Troy McMullin 12 Joel A. Mercado-Daz 13 Matthew P. Nelsen 1, 14 Eimy Rivas Plata 1, 15 William Safranek 16 William B. Sanders 17 Harold P. Schaefer Jr. 18 and Jean Seavey 19 Lcking, R., Seavey, F., R. S. Common, S. Q. Beeching, O. Breuss, W. R. Buck, L. Crane, M. Hodges, B. P. Hodkinson, E. Lay, J. C. Lendemer, R. T. McMullin, J. A. Mercado-Daz, M. P. Nelsen, E. Rivas Plata, W. Safranek, W. B. Sanders, H. P. Schaefer Jr. & J. Seavey. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18 th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127186. 1 Department of Botany, The Field Museum, 1400 South Lake Shore Drive, Chicago, IL 60605 USA; 2 Dan Beard Research Building, Everglades National Park, Homestead, FL 33030 USA < natureguides@mindsping.com>; 3 534 Fenton St., Lansing MI 48910 USA < common@msu.edu>; 4 301 Connecticut Ave., Atlanta, GA 30307 USA ; 5 Naturhistorisches Museum Wien, Botanische Abteilung, Burgring 7, 1010 Wien, Austria ; 6 Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458 USA ; 7 Illinois Natural History Museum, Natural Resources Bldg., 607 E. Peabody Dr., Champaign, IL 61820 USA < leecrane@inhs.uiuc.edu>; 8 The Nature Conservancy in Georgia, 1330 West Peachtree Street, Suite 410, Atlanta, GA 30309 USA < mhodges@tnc.org>; 9 Department of Biology, Box 90338, Duke University, Durham, NC 27708, USA < brendan.hodkinson@duke.edu>; 10 239 Marlborough St., Boston, MA 02116, USA ; 11 Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY 10458, USA ; 12 Biodiversity Institute of Ontario Herbarium, Department of Integrative Biology, University of Guelph, Guelph, ON N1G 2W1, Canada ; 13 International Institute of Tropical Forestry (USFS), Jardn Botnico Sur, 1201 Calle Ceiba, San Juan, PR 00926, Puerto Rico ; 14 Committee on Evolutionary Biology, University of Chicago, 1025 E. 57 th Street, Chicago, IL 60637 USA ; 15 Department of Biological Sciences, University of Illinois-Chicago, 845 West Taylor Street (MC 16 Department of Molecular Biology and Microbiology, College of Medicine, University of Central Florida, Orlando, FL 32816, USA ; 17 Department of Biological Sciences, Florida Gulf Coast University, Ft. Myers, FL 33965, USA < wsanders@fgcu.edu>; 18 25 A Medway Street, Dorchester, MA 02124, USA ; 19 Dan Beard Research Building, Everglades National Park, Homestead, FL 33030, USA ; 20 are in alphabetical order.

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128 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Lcking, (Nyl.) Lcking & Rivas Plata, and (Zahlbr.) Lcking Nyl., Nyl., (Hale) Mangold, Mont. & Bosch, A. L. Sm., and Makhija & Patw. Further 83 species are additions to the North American lichen checklist: (Malme) Marbach, (Nyl.) R. C. Harris, (Fe) Nyl., Vain., (Malme) R. Sant., Lcking, Mll. Arg., (Vain.) Zahlbr., Calopadia (Nyl.) Rivas Plata & Mangold, Chapsa (Tuck.) Breuss & Lcking, C. W. Dodge, (P. Henn) Lcking, Rivas Plata, Lcking & Umaa., G. Thor) Rivas Plata, Lcking, Umana & Chaves., Lcking, Aptroot & Sipman., (Mll. Arg.) Grube, (Tuck.) D. Hawksw. & Dibben, (Mll. Arg.) R. Sant., Sparrius., (Mll. Arg.) Zahlbr., f. Parm., (Mll. Arg.) R. Sant., (Tuck.) Lcking, Srus. & Kalb, (Nyl.) Staiger, (Nyl.) Nyl., (Zahlbr.) Lcking & Rivas Plata, Mont., (Nyl.) Staiger, (Mll. Arg.) Lcking, Lcking & Umaa., Nyl., Redinger, Lcking, Zenker., Vain. Lcking, Nyl., Zahlbr., Kremp., M. Nakan., Lcking, Leight., M. Cceres & Lcking, Lcking & Chaves., Mll. Arg., Herrera-Campos & Lcking, Lcking & G. Thor, Aptroot, Lcking & Will-Wolf., Nyl., Stizenb., (Nyl.) A. Frisch., (Mll. Arg.) Kalb & Lcking, (Vain.) Kalb & Lcking, (Mll. Arg.) Kalb & Lcking, (Spreng.) Kalb, Rivas Plata & Lumbsch, (Tuck.) Kalb, Rivas Plata & Lumbsch, Kalb, (Eschw.) Kalb, Rivas Plata & Lumbsch, (Nyl.) Hale, (Nyl.) Mll. Arg., (Nyl.) Hale, Dibben, Dal Forno & Eliasaro., (Fe) Mll. Arg., (Kremp.) Mll. Arg., (Kremp.) Lcking, (Vain.) Zahlbr., (Ach.) Staiger, (Mll. Arg.) Mll. Arg., Timdal, Mll. Arg., Mll. Arg., Henssen, Fe, (Nyl.) Mll. Arg., Lcking & Lumbsch, Fr., R. Sant., Lcking, R. Sant., and Nyl. The high number of species found within a relatively small area, which corresponds to almost 10% of all lichens currently known in North America, is put into perspective by comparing it with other protected areas in the United States. It is explained by the high carrying capacity of (sub-)tropical vegetation for epiphytic and particularly crustose lichens. Key Words: Ascomycota; lichens; new species; biodiversity; biogeography; Florida

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 129 INTRODUCTION Lichens have long been believed to be most abundant and speciose in cool-temperate areas, a notion supported by a recently published survey from Alaska and its global comparison with other published inventories (Spribille et al. 2010). compare areas of very different size (of more than two orders of magnitude) and also include heterogeneous accounts of lichenicolous fungi. At smaller scales, e.g. looking at uniform area sizes of 100 km 2 10 km 2 1 km 2 or even one hectare, lichen species richness clearly increases towards lower latitudes, like that of many other organisms, a fact realized only recently. This is particularly true for crustose epiphytic microlichens which, for many groups, have their highest diversity in the tropics (Aptroot 1997; Aptroot & Sipman 1997; Sipman & Aptroot 2001; Feuerer & Hawksworth 2007; Lcking et al. 2009). Lcking et al. (2009) estimated that about half of the predicted global 26,000,000 lichen species occur in the tropics, even considering that the tropics cover less than 25% of the Earths land surface. Our ongoing surveys in the Neotropics, particularly in Mexico (Los Tuxtlas Biosphere Reserve), Costa Rica (Las Cruces and La Selva Biological Stations, Corcovado National Park), Venezuela (Henry Pittier National Park), and Peru (Los Amigos Biological Station) suggest numbers of 500 or more species per km 2 of rain forest vegetation, about half of them foliicolous (Lcking 1999; HerreraCampos et al. 2004; Lcking, unpubl. data). In quantitative studies, inventories revealed totals of TABLE OF CONTENTS Introduction .......................................................................................................... 129 Materials and Methods ......................................................................................... 132 Results and Discussion ........................................................................................ 137 New or Otherwise Interesting Species ................................................................. 138 Hodges & Lcking, new species .................................. 138 Calopadia imshaugii Common & Lcking, new species .............................. 139 (Tuck.) Breuss & Lcking, new combination .......... 140 Vain. ex Lcking, new species .................................... 140 Common & Lcking, new species ............................ 141 Common & Lcking, new species ......................... 142 Common & Lcking, new species ................................... 142 Common & Lcking, new species .................................. 143 (Zahlbr.) Lcking & Rivas Plata, new combination (with remarks on (Nyl.) Lcking & Rivas Plata, new combination) ........................................................................... 144 Lcking & Rivas Plata, new species ................ 145 Common & Lcking, new species ........................ 145 Common & Lcking, new species ............................ 146 Common & Lcking, new species ........................... 147 Common & Lcking, new species ......................... 148 Common & Lcking, new species .............................. 149 Lcking & Rivas Plata, new species ........................... 149 Annotated Checklist of Lichen Species from Fakahatchee Strand Preserve State Park ........................................................................................ 150 Acknowledgements .............................................................................................. 178 Literature Cited .................................................................................................... 178

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130 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) approximately 150 corticolous lichen species on 50 trees in the forest understory (Cceres et al. 2008), as many as 250 corticolous species on less than ten trees in the forest canopy (Komposch & Hafellner 1999, 2000), as many as 173 corticolous species on a single tree from top to bottom (Aptroot 1997) and as many as 300 foliicolous species on 100 trees and shrubs (Lcking 1999); as many as 49 foliicolous species were found on a single leaf (Lcking & Matzer 2001). Extratropical sites with similarly high or higher species numbers (excluding lichenicolous fungi) listed in Spribille et al. (2010) are either very large in size, such as Cvennes National Park (France) with 973 lichen species (2297 km 2 ), the Bavarian Forest (Germany) with 841 species (over 9000 km 2 ), Pechoro-Ilychiskiy Nature Reserve (Russia) with 790 species (7213 km 2 ), the Gurktaler Alps (Austria) with 774 species (2950 km 2 ), the dAosta Valley (Italy) with 631 species (3262 km 2 ), and Isle Royale National Park (U.S.A.) with 596 species (2314 km 2 ), or include a heterogeneous and diverse geography, such as Tatra National Park (Poland) with 864 species (211 km 2 ), Berchtesgaden National Park (Germany) with 813 species (210 km 2 ), Klondike Gold Rush National History Park (U.S.A.) with 668 species (53 km 2 ), and Vega Island (Norway) with 649 species (163 km 2 ). None of these areas maintain such large numbers at the small scale, but rather they are the result of combining different vegetation and substrate types with substantial beta for the Klondike Gold Rush National History Park (Spribille et al. 2010) is composed of 668 lichen species and 98 lichenicolous fungi, which were sampled from two different units within the park. The larger unit, Chilkoot Trail, harbored 510 lichen species within an area of 35 km 2 Considering that these include lichens from all substrata (rock, soil, bark, bryophytes), that number is substantially lower than 500 species of lichens that can be found solely epiphytic within 1 km 2 of tropical rain forest. The high species richness in the tropics is attributed to several different factors (Shmida & Wilson 1985; Stocker et al. 1985; Linsenmair 1990), including the absence of frost and the generally nearoptimal conditions for photosynthesis, the strategy of escaping competition by being rare and having effective reproductive mechanisms, a tendency for tropical communities to have a high level of entropy or disorder by dispersion of populations, high dynamics with frequent intermediate disturbances (Connell 1978; Huston 1979; Schupp 1992; Burkey 1994; Hubbell et al. 1999) which, along with biological feed-back mechnisms (Janzen 1970; Becker et al. 1985; Pirozynski & Hawksworth 1988; Armstrong 1989; Waterman & McKey 1989; Wills et al. 1997) prevent competitive species from becoming dominant and instead create a mosaic of successional stages within communities, and the notion that diversity begets diversity in an autocatalytic fashion. Some of these hypotheses have also been tested in tropical lichens (Lcking & Bernecker-Lcking 2000, 2002; Lcking 2008). The continental United States covers an area ranging from the Arctic in the north to the subtropical deserts and subtropical coastal swamp forests in the south. Accordingly, the number of lichen species estimated to occur within this area is large, possibly close to 5,000 species. Most areas and ecosystems are comparatively well-studied with regard to their lichen biota (Harris 1990, 1995; Bennett & Wetmore 1999, 2005a, b; Brodo et al. 2001; Bennett 2006; Thomson 2003; Esslinger Greater Sonoran Desert Region, with about 1,800 species included in a recent monographic treatment (Nash et al. 2002, 2004, 2007). Other lichen-rich regions that have been partially inventoried are the Ozarks and part of the Appalachians in eastern North America (e.g. Hale 1957; Dey 1978; Peck et al. 2004; Harris & Ladd 2007, 2008; Knudsen et al. 2007; Amtoft et al. 2008; Harris 2009; Harris & Lendemer 2009; Lendemer 2009a, b, c). America are mostly restricted to the peninsula of Florida extending westward along the coast through Alabama, Mississippi, Louisiana, and Texas. But some usually tropical elements that can tolerate more temperate conditions, including

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 131 some foliicolous lichens, also extend northward into coastal Georgia and the Carolinas and in some cases the Delmarva Peninsula and southern New Jersey (Lcking et al. 2007; Lendemer & Knapp 2007; Lendemer & Yahr 2004). Recently disjunct populations of typically tropical taxa have also been uncovered at middle and low elelvations of the southern Appalachian Mountains (Lendemer 2009; Lendemer & Tripp 2008, and unpubl. data). Many treatments on tropical lichens are available for these areas (Moore 1966, 1968; Skorepa 1968; Reese & Tucker 1970; Srusiaux 1979; Tucker 1979, 1981; Tucker & Harris 1980; Smith 1986; Harris 1987, 1988, 1990, 1995; 1995; Lumbsch et al. 1995; Ekman 1996; Staiger & Kalb 1999; Marbach 2000; Buck & Srusiaux 2000; Grube 2001), including the classic works by Calkins (1885, 1886, 1889; Eckfeldt & Calkins 1887a, b), Mller Argoviensis (1895), Rolfs (1901), Merrill (1913), Bouly de Lesdain (1933), and Herre (1942). Despite the large amount of lichen diversity documented in these works, which span over a century, previously unknown and unreported taxa continue to be found in the region at a remarkable rate (Lcking & Cceres 2004; Safranek & Lcking 2005; Perlmutter 2006; DeBolt et al. 2007; Lcking et al. 2007; Lendemer & Knudsen 2008; Lendemer & Lumbsch 2008; Grube & Lendemer 2009; Lendemer et al. 2009a d; Seavey 2009; Seavey & Seavey 2009; Tucker 2010; Nelsen et al. 2010; Tripp et al. 2010). These papers document the extraordinary diversity of the regions lichens, especially of crustose epiphytic microlichens, suggesting that many more species await discovery and could provide substantial additions to the North American Lichen Checklist (Esslinger 2010). With this in mind, the 2009 Tuckerman workshop (Fig. 1), as part of the annual lichen workshops organized by the Eastern Lichen Network (http://www.nybg. org/bsci/lichens/eln) was held at Fakahatchee Strand Preserve State Park, which together with Everglades National Park, Big Cypress National Preserve and Collier-Seminole State Park, form a large protected area covering the southern and Figure 1. Group photograph of the 2009 Tuckerman workshop. Standing from left to right: Lee Crane, Sean Beeching, Frederick Seavey, Malcolm Hodges, Richard Fagan, William Buck, Elisabeth Lay, Mike Owen, Harold Schaefer, Matthew Nelsen, William Sanders, Troy McMullin, and Brendan Hodkinson. Kneeling from left to right: Jean Seavey, Bob Nesmith, Alicia Campanella, James Lendemer, William Safranek, Ottmar Breuss, Joel Mercado-Daz, Eimy

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132 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) southwestern part of Florida peninsula. The workshop was organized as part of an ongoing NSF workshop project towards an inventory of neotropical crustose epiphytic microlichens (Lcking et al. 2009; http:// www.nsf.gov/awardsearch/showAward. do?AwardNumber=0715660) and served several objectives: extending the area covered by the annual Tuckerman workshops to southern Florida and thus the most tropical part of the continental United States; document in more detail the tropical element of the lichen biota of the continental United States; train participating colleagues and students in tropical lichen taxonomy, with emphasis on crustose epiphytic microlichens; and produce a guide to crustose epiphytic microlichens found in the area to be used by the park personnel and ecotourists. MATERIALS AND METHODS STUDY AREA Fakahatchee Strand Preserve State Park, a narrow strip of land oriented north to south, is located in Collier County, Florida, and is about 3 miles (4.5 km) wide and 20 miles (36 km) long (Fig. 2). Hydrologically, it is a shallow and narrow drainage basin channeling the waters from low-lying terrains to the north and east into the Ten Thousand Island zone of the Gulf of Mexico (Petuch & Roberts 2007). The name Fakahatchee is derived from the Seminole Indian language which, roughly translated, means muddy creek. During the summer rainy season from May to October, which provides about 42 of the annual 53 inches of precipitation, most of the Preserve is inundated to varying depths. This is reversed in winter months Figure 2. A, County map of Florida showing Collier County and the study site. B, Map of Fakahatchee Strand Preserve State Park, straddling the Tamiami Trail (Route 41) within Collier County; the four main collection sites described in the text are indicated by numerals 1 to 4. Map reprinted with permission of the Florida

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 133 as ground water recedes and in drier years much of the Preserve may lack surface water. Winter temperatures are mild, ranging normally from 10C to 18C. The mean January low temperature is slightly below 12 which, in the outside the subtropical zone. Occasional cold fronts may bring mild frosts. Summer months are very humid with overnight lows rarely dropping below 18C and daytime highs around 32C. The rock substrate is predominately of biogenetic limestone (Petuch & Roberts 2007) but outcrops projecting lichen colonization are rare. Although logged extensively during the 1940s and 1950s, relatively mature secondary forests now cover a large portion of the Preserve. In the north and central part, the major tree species include (bald cypress), (dahoon), (Carolina ash), (royal palm), (cabbage palm), (red maple) and (live oak). The mangrove zone to the south supports (red mangrove), (black mangrove), (white mangrove) and, on higher ground, (buttonwood). Situated between a West Indian-dominated for its vascular plant diversity by sharing both Encompassing only a little over 85,000 acres (34,000 hectares), at least 524 vascular plant species have been recorded from the Preserve (Institute for Regional Conservation online database: http:// regionalconservation.org/ircs/reports_database. asp ). Outside South Florida, many of these are found nowhere else in North America and some are endemic to the park (Avery & Loope 1980; Austin et al. 1990). By comparison, Everglades National Park, with its northwest boundary located 1,200 plant species, although it occupies nearly 1.4 million acres (560,000 hectares) (Olmstead & Loope 1984; Avery & Loope 1996). has been effectively inventoried, the lichen biota has not. Due to its climate and geological history, southern Florida, including the Preserve, possesses United States (Beard 1938; Robertson 1955; Olmstead et al. 1981). It is then logical to assume the lichen biota should also be different. The most extensive lichen collections from the Preserve were made in 1992 by Richard Harris and William Buck and in 1997 by Ralph Common. These forays produced collections of several hundred species. A few collections were also taken by Sylvia and Stephen Sharnoff while accumulating photos for the book coauthored with Dr. Irwin Brodo in 2001 (Brodo et al. 2001). Undoubtedly, other unpublished and unrecorded collections have been made but, if maintained at all, they appear to be scattered. Currently, no single database or inventory dedicated to the lichens of the Preserve is maintained (R. Rau, Park Manager at Fakahatchee Strand Park Preserve, pers. comm. 2009). The collections previously made by others plus those gathered during the 2009 Tuckerman Workshop now document over 400 species and can be considered representative. This publication should provide the Preserve with the framework on which to build a lichen checklist which will, we hope, be as comprehensive as that of vascular plants. COLLECTION SITES Although small in relation to its giant neighbors Big Cypress National Preserve and Everglades National Park, the Preserve contains many distinct ecological communities including mangrove forests, prairies of grasses and sedges, nearly monotypic cypress stands and a variety of broad leaf forest types. However, due to the limited collecting time allotted (two full days in addition to time assigned for laboratory work), only four sites were selected for sampling. The chosen sites featured ease of access, a close proximity to one At Site 1 (Figs. 2B, 3 AB ), a lowland forest is dominated by a unique canopy mix of royal

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134 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) palms and bald cypress. Dahoon and Carolina ash are common in the subcanopy while the understory varies from dense to open. In most places at least features a raised tram road once used by logging companies to extract cut trees (Fig. 3 CD ). Lichens are abundant along the margins of the road, particularly foliicolous species on cabbage and saw palm leaves ( ). Collectors also wandered off into the short canopy forests on either side of the road. Species of and are common at this site. Site 3 straddles the main road through bald cypress stand on the east side of the road and an upland hammock (tree island) on the west. The latter is dominated by live oak, dahoon, cypress and red maple (Fig. 3EF) Judging by the large number of bald cypress stumps and cypress knees, Site 4 was once a cypress swamp forest (Fig. 3G H) Although some cypress has regenerated since logging days, the site today is dominated by dahoon, Carolina ash, red maple, and swamp dogwood ( ). The understory was open during the lower strata of the community. During summer months, site 4 is inundated to various depths but at the time of collecting ground water was absent except for a few scattered ponds. SPECIES IDENTIFICA TIONS Specimens were studied using standard techniques of light microscopy and thin-layer Edison State College in Naples, Florida and subsequently at the institutions of the participants. The information was compiled by Rick and Jean Seavey and Robert Lcking and each reported taxon was critically revised to ensure were compared with authentic type material, particularly for the Graphidaceae (including Thelotremataceae), Porinaceae, Pyrenulaceae, and Trypetheliaceae. Specimens were studied using a diverse array of optical equipment at Edison State College and subsequently at the institutions of each participant. For spot reactions and microscopic staining, we used 10% potassium hydroxide solution (K), commercial bleach containing 6% sodium hypochloride (CHLOROX), paraphenyldiamine crystals dissolved in 95% ethanol, and Lugol solution (FLUKA 62650). Standard thin-layer chromatography (TLC; Arup et al. 1993; Lumbsch 2002) was applied to a large number elucidate more complex chemical patterns. Most images were taken from freshly collected material at Edison State College using a NIKON Coolpix 5400 digital camera mounted on one ocular of a stereomicroscope. SPECIES RICHNESS In order to put the number of lichen species found here into a wider context, we compared the data to those published for other nature preserves in the continental United States (Bennett & Wetmore 2005a; Spribille et al. 2010) and extracted the most recent data of the NPLichen database for national parks and other nature preserves (Bennett & Wetmore 2005b, Bennett 2006; http://www.nbii. gov/portal/server.pt/community/lichens/1555 ). We also used these authors estimates of relative completeness of the lichen inventories in these parks and preserves to account for missing data, knowing that this can only be an approximation and that many parks require more detailed studies to assess their lichen diversity accurately. It should be noted that recent surveys of several parks considered 90% known in the NPLichen database (e.g. the Santa Monica Mountains and the Great Smoky Mountains; Knudsen & 2010; Lendemer et al., unpubl. data) exhibit undiscovered lichen species richness that was higher than previously estimated. However, this applies likely to other parks and preserves, including Fakahatchee, and therefore the level of uncertainty is partially balanced out and the relative comparisons remain valid at this point. Assuming that lichen species richness depends more or less logarithmically on area and rather linearly on habitat diversity (McGuiness 1984; Nilsson et al. 1988; Rosenzweig 1995; NeyLcking et al. 2009a), in our analysis we used the

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 135 Figure 3. Habitat photographs of the four collection sites at Fakahatchee Strand Preserve State Park. First row: site 1, showing royal palm canopy and understory with bald cypress. Second row: site 2, showing side road along old tram with dense vegetation and aspect of understory with bromeliads. Third row: site 3, showing open bald cypress prairie hammock and dense regrowth. Fourth row: site 4, showing closed bald cypress forest and aspect of forest understory. Photographs by R. Lcking (A, B, G),

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136 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Table 1. Lichen species richness of selected North American national parks and other preserves (after Bennett & Wetmore 2005a, b; Bennett 2006; Spribille et al. 2010; http://www.nbii.gov/portal/server.pt/community/lichens/1555 ). SPE = currently known species richness; COMP = estimate of completeness range; EST = estimated species richness using mean from estimate of completeness range (mean value; the range would be 5%); SIZE = area cover (excluding water surfaces); ALT 1998); SPE/area = species per area; SPE/log = species per log-area; SPE/veg = species per log-area per vegetation categories. For further explanation see text. *For these parks, species versus log-area relationships have a steeper slope due to small area size and hence comparison with larger areas does not apply and SPE/area, SPE/log, and SPE/veg calculations are not given. **Figures updated according to list maintained by park; further 90 species are to be added and hence the lichen species richness of that park will surpass Kondike (Lendemer, pers. comm. 2010). PARK NAME SPE COMP EST SIZE [km 2 ] ALT [m] VEG SPE/ area SPE/ log SPE/ veg Klondike (Alaska)* 668 91% 703 53 Isle Royale (Michigan) 613 91% 645 540 180 6 1.14 2.24 0.37 Great Smoky Mountains (North Carolina)** 589 91% 620 2100 265 7 0.28 1.77 0.30 Voyageurs (Minnesota) 498 91% 525 250 335 5 1.99 2.08 0.42 Glacier (Montana) 480 91% 505 3830 960 7 0.13 1.34 0.19 Everglades (Florida) 450 76% 540 6000 0 5 0.08 1.19 0.24 Acadia (Maine)* 444 91% 465 140 Gates of the Arctic (Alaska) 441 91% 465 39000 85 6 0.01 0.96 0.16 Fakahatchee Strand (Florida) 432 76% 519 320 0 5 1.35 1.72 0.43 Yellowstone (Wyoming) 421 76% 505 8500 1610 7 0.05 1.07 0.15 Rocky Mountain (Colorado) 419 91% 440 1050 2325 6 0.40 1.39 0.23 Keweenaw (Michigan)* 354 91% 375 8 Olympic (Washington) 353 76% 425 3700 0 7 0.10 0.99 0.14 Delaware Water Gap (New Jersey) 333 91% 350 265 90 6 1.26 1.37 0.23 Santa Monica Mountains (California) 333 91% 350 620 0 7 0.54 1.19 0.17 Chiricahua (Arizona)* 325 91% 340 120 Apostle Islands (Wisconsin) 323 91% 340 160 N/A 5 2.02 1.47 0.29 Saint Croix (Minnesota) 308 91% 325 350 N/A 5 0.88 1.21 0.24 Saguaro (Arizona) 299 91% 315 370 665 7 0.81 1.16 0.17 Big Bend (Texas) 289 91% 305 3200 520 8 0.09 0.82 0.10 Channel Islands (California) 273 91% 285 500 0 7 0.55 1.01 0.14 Pictured Rocks (Michigan) 263 91% 275 290 N/A 5 0.91 1.07 0.21 Denali (Alaska) 259 51% 410 20183 70 6 0.01 0.60 0.10 Sequoia (California) 256 91% 270 1600 415 9 0.16 0.80 0.09 Grand Canyon (Arizona) 255 51% 405 4900 780 7 0.05 0.69 0.10 Theodore Roosevelt (North Dakota) 251 91% 265 280 590 5 0.90 1.03 0.21 Redwood (California) 251 76% 300 450 N/A 7 0.56 0.95 0.14 Blue Ridge (North Carolina/Virginia) 246 51% 390 370 200 8 0.66 0.96 0.12 Mount Rushmore (South Dakota)* 222 76% 265 5 Grand Teton (Wyoming) 221 51% 350 1250 1900 7 0.18 0.71 0.10 Shenandoah (Virginia) 218 76% 265 800 180 6 0.27 0.75 0.13 Hot Springs (Arkansas)* 214 91% 225 22 Bandelier (New Mexico)* 211 91% 220 130 Costa Rica Las Cruces 550 91% 580 2.7 1200 1 203.70 12.75 12.75 Costa Rica La Selva 598 91% 630 28 200 1 21.36 4.13 4.13 Costa Rica Tortuguero 523 76% 630 190 0 1 2.75 2.30 2.30 Costa Rica Hitoy Cerere 485 76% 585 200 200 1 2.43 2.11 2.11 Costa Rica Corcovado 511 76% 615 420 0 1 1.22 1.95 1.95

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 137 following parameters: total number of species, species per area, species per log-area, and species per log-area per habitats. For each preserve, the total area was corrected to the area covered by land. To assess the number of principal vegetation types (suitable for lichen growth) in each preserve, Grossman et al. (1998). In addition to principal vegetation types, the number of altitudinal belts was taken into consideration, using a relative assessment depending on latitude, since the tree line and hence the potential substrate for epiphytic lichens decreases with higher latitudes. RESULTS AND DISCUSSION Together with previous collections gathered at Fakahatchee Strand Preserve State Park by R. Common, W. R. Buck, and R. C. Harris, the total number of species currently documented for the Preserve amounts to 432, most of which were collected during the 2009 Tuckerman workshop. All species are listed below, together with representative voucher specimens, and most of the crustose epiphytic lichens are documented with color images. The 432 species represent over 100 genera in 35 families and approximately 2,000 collections in total. Fourteen species new to science are described below (plus four others in three separate papers; Nelsen et al. 2010; Lumbsch et al. 2011; Lcking et al. 2011) and we also report 89 species new to the North American Lichen Checklist (Esslinger 2010). Although South Florida lies entirely north of similar to those of the Caribbean (Tomlinson 1980). Tomlinson also stated that the 12C January mean temperature isotherm for the state was most of Florida between the shoreline and this isotherm would be where the concentration of Caribbean species would be found. With minor exceptions, this has proved to be the case (Olmstead et al. 1993). Fakahatchee Strand Preserve State Park narrowly misses being within this zone, but still has (mastic), (satinleaf) and (indigoberry). Conversely, several arboreal species common at Fakahatchee apparently have reached their southern limits and become scarce or absent only a few miles to the south of the site. These include (Carolina ash), (red maple), and (laurel oak). The complete botanical checklist of the Preserve reveals essentially the same type of distributional composition and demonstrates that it is a geographical ecotone between temperate and tropical biomes. The 432 lichen species found at Fakahatchee show a similar mixture of temperate and tropical taxa, with a distinct bias towards tropical species, and this certainly accounts for the richness of the preserves lichen biota. The majority of the species recorded here as new to the North American lichen including and have been known so far only from the Old World tropics. Others are essentially tropical but have been collected beyond tropical latitudes under favorable conditions, such as foliicolous species (Lendemer & Tripp 2008). The high percentage of species found to be new to North America within the preserves newly created checklist can in part be attributed to the paucity of previous systematic lichen explorations at this location, as only few collections have been studied and published before (Harris 1990, 1995). Several lichenologists have collected at Fakahatchee but there is no record of any lichen foray equaling the present one. The list below also includes some taxa that keyed imperfectly or would not key at all and require further study, most of them lacking ascomata. The checklist consists mostly of corticolous and foliicolous collections with only a few from lignum or on moss over bark. The lack of saxicolous specimens is accounted for by the nature of the geology of the collection site and South Florida in general. For many million years the area has been formed by succesive layers of biogenic marine carbonates, mostly limestone (Randazzo & Jones 1997; Petuch & Roberts 2007). It has been

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138 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) covered by peat and calcareous marls following its emergence above sea level early in the Pleistocene. Although limestone outcrops are common, many of these are inundated nearly half the year by summer rains. Siliceous rocks are absent from the region. For the most part, only the higher prairies (Orzell & Bridges 2006) contain exposed calcareous outcrops potentially capable of was not sampled during this foray. Although Fakahatchee is a small park, being 22 nd in size of all parks with more than 200 lichen species reported and compared here (Table 1) and only 10% of the average size of these parks, i t currently ranks 9 th in lichen species richness. By far the largest numbers of species have been documented for Klondike Gold Rush National History Park and Isle Royale National Park, with 668 and 613 taxa, respectively (Table 1). Several other parks have between 400 and 500 species, including Everglades National Park (Seavey & Seavey, unpubl. data). In addition to the parks and preserves listed in Table 1, further inventories are currently under way. Tucker (2010) recently reported 296 lichen species from Burden Research Plantation in Louisiana, another subtropical site that has been monitored for over 25 years. Hodkinson (2010) and Hodkinson et al. (2009, 2010) found 220 lichen species at Mount Rogers National Recreation Area. Excluding parks of less than 200 km 2 (because the log-area approximation does not directly compare between small and large area sizes due to changes in the slope parameter; McGuiness 1984; Nilsson et al. 1988; Rosenzweig Strand Preserve State Park ranges among the top species per log-area per habitat indices (Table 1). The latter index surpasses that of much larger parks with comparable vegetation, such as the nearly twenty times larger Everglades National Park, and underlines the carrying capacity of (sub-)tropical vegetation to support high small-scale diversity, especially for epiphytic crustose microlichens. The North American lichen checklist currently comprises more than 4500 lichen species (Esslinger 2010), which means that Fakahatchee supports almost 10% of the lichen species richness reported for the entire North America, although it is only a fraction of its size (0.00015%). This corresponds to a 60000-fold increase in species density. This phenomenon of concentrated, high small-scale species richness comes to its extreme in the tropics, such as in Costa Rica, where the species per log-area in any species-rich park found in the continental Unites States, including the (sub-)tropical parts of Florida (Table 1). NEW OR OTHERWISE INTERESTING SPECIES In the following account, we present taxonomic novelties found during this and previous lichen surveys at Fakahatchee. Other new species are described in separate papers (Nelsen et al. 2010; Lumbsch et al. 2011; Lcking et al. 2011). The taxa are listed in alphabetical order. Hodges & Lcking, new species Figure 14DG 1 Mycobank #518000 Diagnosis.Sicut apotheciis pruinosis maioribusque differt. Description.Thallus corticolous, 1 cm diam., 30 m thick, centrally continuous but towards the margin with dispersed, irregular patches; surface uneven, white to pale gray. Photobiont chlorococcoid. Apothecia sessile, rounded, 0.6 1.2 mm diam. and 250 m high; disc plane to slightly convex, dark gray to brownish gray, coarsely white-pruinose; margin thin, pale gray to cream-colored. Excipulum paraplectenchymatous, 50 m broad, colorless. Hypothecium 30 m high, dark brown. Apothecial base aeruginous. Epithecium thin, 5 m high, pale grayish 1 Figures 4-58 are available on-line at http://www.

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 139 brown. Hymenium 120 m high, colorless. Paraphyses unbranched to slightly branched. Asci 100 25 m. Ascospores single, ellipsoid, muriform, 50 17 m, 3 times as long as broad, colorless. Campylidia sessile, 0.6.2 mm broad, 1.5 mm long; lobe well-developed, hoodshaped, dark gray with paler apex, white-pruinose; apex, 5-septate, 50 1.5 m. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Janes Scenic Drive 6.5 mi NNW of ranger station, west of old tram, hardwood hammock, slough and strand, on branch, March 2009, (F, holotype). Etymology.Referring to its discovery in Florida. Distribution and Ecology.This species is known from two well-developed collections from Fakahatchee Strand Preserve State Park and Sumpter County, growing on the smooth part of the bark of small branches and partly on exposed areas of the wood bare of periderm, as well as on hardwood bark. Remarks. belongs in a group of species that have single, muriform ascospores less than 100 m long. All species so far known in this group, except the recently described (see below in the species list), have non-pruinose apothecia which are also distinctly smaller, usually not exceeding 0.8 mm diam. also has pruinose apothecia but the disc is lighter brown and yellowish pruinose and the pruina also covers the apothecial margin. Except for the pruina on the apothecial disc, the new species comes closest to also present in the park, which shares the dark brown hypothecium and aeruginous apothecial base and a gray-brown apothecial disc, but differs in its smaller, non-pruinose apothecia. Another species with pruinose apothecia and single-spored asci is which has longer ascospores (> 100 m) and the apothecia, as in are yellow-brown with pale yellow-white pruina (Lcking 2008). The holotype of bears only mature apothecia, all heavily pruinose, whereas an additional collection from Sumpter County ( ) has younger apothecia with scarce pruina only, suggesting that the pruina develops with age. Additional Specimens Examined.U.S.A. Florida. Sumpter Co.: Rt. 471 along Withlacooche River, at boundary with Polk Co., Apr 1992, (hb. Common). Calopadia imshaugii Common & Lcking, new species Figure 15AF Mycobank #560000 Diagnosis.Sicut sed apotheciis maioribus pruinosis et ascosporis 1 in ascis differt. Description.Thallus corticolous, 0.5 cm diam., 30 m thick, centrally continuous but towards the margin with dispersed, irregular patches; surface uneven, white to pale gray. Photobiont chlorococcoid. Apothecia sessile, rounded, 0.4.8 mm diam. and 200 m high; disc plane to slightly convex, brown, coarsely white-pruinose; margin conspicuous, pale gray to cream-colored. Excipulum paraplectenchymatous, 50 m broad, colorless. Hypothecium 20 m high, (dark) brown. Apothecial base sordid brown. Epithecium thin, 5 m high, pale grayish brown. Hymenium 100 m high, colorless. Paraphyses unbranched to slightly branched. Asci 90 20 m. Ascospores (1)2() per ascus, ellipsoid, muriform, 50 15 m, 3 times as long as broad, colorless. Campylidia sessile, 0.5 mm broad, 1.5 mm long; lobe well-developed, hood-shaped, dark gray with paler apex, white-pruinose; socle not apparent. Conidia 1.5 m. Secondary chemistry: all spot tests negative. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.We dedicate this new species

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140 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) to the late Henry Imshaug for his contributions to lichenology. Distribution and Ecology.This species is known from several collections in Florida, growing Remarks. Calopadia imshaugii is unusual within the genus in having more than one ellipsoid ascospore per ascus. Most species of the genus have single-spored asci, and in the few taxa with more than one ascospore per ascus, the ascospores are narrow and oblong rather than ellipsoid, being 70 8 m (Lcking 2008). The ascospores of the new species have the same shape and dimensions as those in species with single-spored asci related to Morphologically, the new species is similar to (described above), but the latter has single-spored asci. Additional Specimens Examined. U.S.A. Florida. Dade Co.: SW 388th St., 1.2 miles east of Old Dixie Hwy., near Homestead, May 1992, (all hb. Common). (Tuck.) Breuss & Lcking, new combination Figure 17D Mycobank #517995 Basionym Tuck., Proc. Am. Acad. Arts Sci. 6:270 (1866); (Tuck.) Mll. Arg., Flora 69:311. (1886). Type.Cuba, (FH!; lectotype, Remarks.This species is closely related to with which it co-occurs in the same microhabitats, mostly on swamp dogwood. The species was originally separated from C. because of the smaller apothecia and larger ascospores with more numerous septa. However, while the two species do differ in apothecial size, the ascospores are practically identical, as already noted by Hale (1981). We found two other distinctive features of Chapsa : the outer thallus margin is more or less entire compared to the distinctly lobulate and recurved margin of and the apothecial margin is UV+ yellow (sometimes only weakly so), indicating the presence of lichexanthone. The species somewhat resembles which also contains stictic acid, but has much longer, multiseptate, hyaline ascospores (Rivas Plata et al. 2010). Additional Specimens Examined: U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Off James Scenic Drive west of Tram Rd. 6.5 mi NNW of Ranger Station, TaxodiumSabal-hardwood hammock, slough, and strand; March 2009, (LI, USF), (F, USF). Vain. Lcking, new species Figure 20EF Mycobank #560001 Diagnosis. speciei medulla cinnabarina differt. Thallus ecorticatus, virido-albus. Ascomata desunt. Acido 2O -methylperlatolico continens. Description.Thallus corticolous, to 10 cm diam., 25 m thick, continuous; surface uneven, minutely farinose-byssoid, pale green to almost white, with mottled areas of pure white, sterile but potentially ascigerous areas; streaks of medulla frequently exposed, bright orange to cinnabar-red. Photobiont trentepohlioid. Thallus in section lacking upper cortex, with irregular algal layer and areas of concentrated crystalline orange pigments. Asci and ascospores not observed. Secondary chemistry: 2O -methylperlatolic acid and unknown substance close to gyrophoric in detected by TLC (paratype); unpigmented thallus parts C, K, P, I+ light blue (especially white areas), pigmented parts K+ dark purple-violet. Type. G uyana Vryheid Plantation, no date or collector given (TUR-Vainio 28367, holotype). Etymology.The epithet refers to the strongly pigmented parts of the thallus, meaning cinnabar-red color (not to be confused with small). Distribution and Ecology.This species is known from the type material from Guyana and from several collections from Fakahatchee Strand

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 141 Preserve State Park, where it was found on bark of and hardwood in shaded situations. Remarks.The material of this taxon found in TUR suggests that Vainio intended to describe this species under the generic name but never published it. It does not seem to have been mentioned in the literature. However, the Florida collections suggest that this is a ascospores are absent. The general habit, the sterile white areas dispersed over the thallus showing a light blue reaction with iodine, and the secondary chemistry including 2O -methylperlatolic acid, support inclusion within However, the pigmented thallus parts are quite unique in the genus. Thus far, pigments appear to be restricted to the related genus (Aptroot et al. 2009), where they usually occur in the hypothallus and prothallus, as well as the isidioid outgrowths. Since the name coined by Vainio was available, we decided to validate this name rather than using a new one; therefore, Vainios material was selected the Florida material which is then available for molecular studies. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, off James Scenic Drive 6.5 mi NNW of ranger station at gate 12 along old tram road, hardwood hammock, strand and swamp, on bark, March 2009, (hb. Lay), (F). Common & Lcking, new species Figure 23E Mycobank #560002 Diagnosis.Sicut sed lirellis minoribus aggregatibusque differt. Description.Thallus corticolous, 1 cm diam., 40 m thick, continuous; surface uneven to verrucose-bullate, yellowish green to olive-brown. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and scattered clusters of crystals. Li rellae densely aggregate in small clusters, straight to curved, unbranched to sparsely branched, erumpent, with lateral thalline margin, 0.3.7 mm long, 0.15.2 mm wide, 0.12.15 mm high; disc concealed; labia conspicuous, thick, yellowish-white; thalline margin olive-brown. Excipulum entire, apically dark brown, 25 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 60 m high, colorless, clear; epithecium granulose, 5 m high, brown. Paraphyses unbranched, glabrous; periphysoids not observed; asci fusiform, 60 10 m. Ascospores 8 per ascus, ellipsoid, transversely 3-septate, 14 7 m (including 1.5 m thick wall), 2 times as long as wide, with comparatively thin septa and angular-rounded lumina, colorless to pale gray-brown, I. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The epithet refers to the small clusters of aggregate lirellae. Distribution and Ecology.This species is known from several small collections from Fakahatchee Strand Preserve State Park, from bark Remarks.This material is characterized by clearly aggregate, as well as 3-septate, non-amyloid ascospores. It comes closest to (Harris 1995; Staiger 2002; Archer 2009), but the lirellae are much more delicate and form small clusters. is another species with aggregate lirellae and 3-septate ascospores (see below), but in that species, the labia are inconspicuous, the lirellae are gaping, and the clusters are much larger and almost pseudostromatic. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second

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142 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) growth area, Apr 1997, (hb. Common), (hb. Common). Common & Lcking, new species Figure 23FG Mycobank #560003 Diagnosis.Sicut sed thallo corticato et ascosporis typo trypethelio differt. longae, 0.1 mm latae. Ascosporae 3-septate, 13 7 m, I. Acidi lichenum desunt. Description.Thallus corticolous, 0.5 cm diam., 30 m thick, continuous; surface smooth to uneven, pale greenish white. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and clusters of crystals. Lirellae straight to curved, unbranched to irregularly branched, margin, 0.3.5 mm long, 0.1 mm wide, 0.1 mm high; disc concealed; labia inconspicuous to slightly bulging, brown along the slit; thalline margin white. Excipulum entire, apically dark brown, 10 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 70 m high, colorless, clear; epithecium granulose, 5 m high, colorless. Paraphyses unbranched, glabrous; periphysoids not observed; asci fusiform, 60 12 m. Ascospores 8 per ascus, ellipsoid, transversely 3-septate, 13 7 m (including 1.5 m thick wall), 2 times as long as wide, with trypethelioid to thin septa and trypethelioid to anglar-rounded lumina, colorless, I. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The name is a word play F. Distribution and Ecology.This species is known from several rich collections, from Fakahatchee Strand Preserve State Park and other parts in Florida. It grows on bark of hardwood. Remarks. belongs in a complex of species with 3-septate, nonamyloid ascospores and lacking lichen substances, centered around Within this complex, the new species is clearly set apart by its extremely small and delicate lirellae, which are the smallest known among all described species. (see below) agrees in the delicate lirellae with apically dark brown excipulum, as well as the non-amyloid ascospores with trypethelioid septa, but has an endoperidermal, ecorticate thallus. its thallus is mostly ecorticate and the ascospores have more rounded lumina and are frequently weakly amyloid. We also observed conspicuously thickened paraphyses in that species but not in F. Additional Specimens Examined. U.S.A. Florida. Baker Co.: Osceola National Forest near Ocean Pond, Dec 1975, , (all hb. Common). Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, , (all hb. Common). Hillsborough Co.: Hillsborough River State Park, Dec 1990, (all hb. Common). CR 581, 3.2 mi S of I-75, Nov 1995, , (all hb. Common). Polk Co.: CR 54, 0.1 mi E of CR 557, Apr 1996, (hb. Common). Common & Lcking, new species Figure 24B Mycobank #560004 Diagnosis.Sicut sed acido psoromico differt. Thallus corticatus. Lirellae labiatae, 0.5 mm longae, 0.2 mm latae. Ascosporae muriformes, 25 12 m, I+ caeruleae. Acido psoromico continens. Description.Thallus corticolous, 1 cm diam., 50 m thick, continuous; surface uneven to verrucose-bullate, yellowish green to

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 143 olive-brown. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and scattered clusters of crystals. Lirellae straight to curved, unbranched to sparsely branched, erumpent to prominent, with lateral thalline margin, 0.5 mm long, 0.15 0.2 mm wide, 0.15.2 mm high; disc partially exposed, gray-pruinose; labia conspicuous, thick, white; thalline margin olive-brown. Excipulum entire, colorless to apically orange-brown, 25 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 80 m high, colorless, clear; epithecium granulose, 5 m high, brown. Paraphyses unbranched, glabrous; periphysoids not observed; asci fusiform, 80 25 m. Ascospores 8 per ascus, ellipsoid, muriform with 5 1 septa, 25 12 m (including 1 m thick wall), 2 times as long as wide, with comparatively thin septa and angular-rounded lumina, colorless, I+ violet-blue. Secondary chemistry: psoromic acid (major), subpsoromic acid (minor), 2'-O-demethylpsoromic acid (major), medulla P+ yellow. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The epithet refers to the fact that this species was previously confused with F. Distribution and Ecology.Known from several collections from Fakahatchee Strand Preserve State Park and also reported from Brazil by Staiger (2002; under ). The species is usually found on branches of hardwood. Remarks.The Florida material agrees well with the description of given by Staiger (2002), including the verrucose-bullate thallus and the erumpent to prominent lirellae with slightly gaping disc and thick, white labia. Staiger (2002) accepts forms with and without psoromic acid in the same species, but we consider this premature until molecular data clarify the situation. In other genera of Graphidaceae, such as s.lat., psoromic acid is accepted as a species-diagnostic character. Since the type of has no substances, we introduce here the name for the psoromicacid containing specimens. All Florida material contains psoromic acid. presently contains four species with muriform (and amyloid) ascospores and psoromic acid chemistry. Among these, (Nyl.) Staiger is set apart by the very small, almost globose ascospores (about 10 10 m), whereas (A. W. Archer) A. W. Archer has more elongate ascospores (about 18 10 m); both have closed, labiate lirellae with white upper surface. The other two species have comparatively large ascospores (about 30 15 m) and differ from each other morphologically: (see below) labia, whereas (A. W. Archer) A. W. Archer has labiate lirellae with concealed disc which are brown-black along the slit due to a weakly carbonized excipulum; in addition, that species differs in having 2-methoxy-psoromic acid, a substance that can be distinguished from psoromic acid only by HPLC (Archer 2001). The new species, also has labiate lirellae but with partially exposed disc and white labia, thus clearly different from both and F. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (hb. Common), (hb. Common), (hb. Common). Common & Lcking, new species Figure 25B Mycobank #560005 Diagnosis.Sicut sed hymenio insperso et acido stictico differt. Thallus longae, 0.1 mm latae. Ascosporae 3-septate, 12 6 m, I. Acido stictico continens.

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144 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Description.Thallus corticolous, 0.5 cm diam., 30 m thick, continuous; surface smooth to uneven, yellowish to pale greenish white. Photobiont trentepohlioid. Thallus in section with partly prosoplectenchymatous, partly irregular and loose upper cortex, irregular algal layer and scattered clusters of crystals. Lirellae straight to curved, unbranched, immersed, 0.5 mm long, 0.1 mm wide, 0.1 mm high; disc partially exposed, dark gray-pruinose; labia inconspicuous; thalline margin yellowish white. Excipulum entire, brown to apically dark brown, 10 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 50 m high, colorless, strongly and densely inspersed (inspersion persistent in K); epithecium granulose, 5 m high, brown. Paraphyses unbranched, glabrous; periphysoids not observed; asci fusiform, 50 10 m. Ascospores 8 per ascus, ellipsoid, transversely 3-septate, 12 18 6 m (including 1.5 m thick wall), 2 times as long as wide, with trypethelioid septa and lumina, colorless, I. Secondary chemistry: stictic acid (major), constictic acid (trace), thallus section Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The name refers to the strongly inspersed hymenium, an unusual character in (Staiger 2002). Distribution and Ecology.This species is known from several rich collections, all from Fakahatchee Strand Preserve State Park, on hardwood. Remarks. is a typical with an unusual character: a strongly inspersed hymenium. The inspersion resembles that of and allies (Lcking 2009), in that it makes the paraphyses, asci, and ascospores barely discernible, but contrary to the latter, in it does not dissolve in K. Thus far, no species of with inspersed hymenium appears to be known (Staiger 2002). Apart from the inspersion and the darkened excipulum, Fink comes closest, as it Staiger (2002), has the same ascospore type, and also produces stictic acid. has a weakly carbonized excipulum and the same ascospore type, but lacks inspersion and its thallus is endoperidermal and completely ecorticate. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (hb. Common); ibid., K2 trail among royal palms, second growth area, Apr 1997, (hb. Common), (hb. Common). (Zahlbr.) Lcking & Rivas Plata, new combination Figure 25C Mycobank #517999 Basionym. Zahlbr., Ann. Mycol. 19:233 (1921); (Zahlbr.) M. Wirth & Hale, Contr. US Natl. Herb. 36:82 (1963). Type.Mexico, (W). with Nyl. (Harris 1995). Staiger (2002) noted that was described as sterile, and indeed, no ascospores are present in the type (indicated on Mason Hales Index Cards at US). The application of this name is therefore uncertain. There are two other species with similar thallus and lirellae morphology: f. Nyl. and (Zahlbr.) Wirth & Hale. Ascospores are present in both types, being 20 m long in and 25 m long in The by Staiger, but differs in the pseudostromatic lirellae and therefore is recombined in : (Nyl.) Lcking & Rivas Plata, new combination and status [Mycobank # 517996 ;

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 145 Basionym. f. Nyl., Bull. Soc. Linn. Normandie, Sr. 2, 2:121 (1868). Type.New Caledonia, (H-NYL material from Fakahatchee, ascospores are mostly Since the type of is from SE Asia (Bengal), we consider it highly unlikely that F. Harris (1995) gave the ascospore size for as 35 18 m. We have not seen ascospores of such a size and they seem to be highly unusual for a ; probably there is a further species belonging to a different genus involved. The collection bears pycnidia with bacillar conidia about 5 1 m in size. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, ). Along James Scenic Drive, 3.2 mi NNW of Ranger Station; -hardwood strand, swamp and slough, Mar 2009, *, (all F). Lcking & Rivas Plata, new species Figure 25D Mycobank #560006 Diagnosis.Sicut sed lirellis immersis albis et ascosporis angustioribus 0.7 mm longae, 0.15.2 mm latae. Ascosporae 3-septate, 14 7 m, I. Acidi lichenum desunt. Description.Thallus corticolous, 1 cm diam., 40 m thick, continuous; surface smooth to uneven, olive-green. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and scattered clusters of crystals. Lirellae densely aggregate in pseudostromatic, white clusters, straight to curved, unbranched, immersed, with thin thalline margin, 0.1.3 mm long, 0.1 mm wide, 0.1 mm high; disc concealed to slightly gaping; labia inconspicuous, white; thalline margin yellowwhite. Excipulum entire, orange-brown, 15 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 60 m high, colorless, clear; epithecium granulose, 5 m high, orange-brown. Paraphyses unbranched, glabrous; periphysoids present, glabrous; asci fusiform, 60 10 m. Ascospores 8 per ascus, ellipsoid, transversely 3septate, 15 5 m (including 1 m thick wall), 3 times as long as wide, with comparatively thin septa and angularrounded lumina, colorless to pale gray-brown, I. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Janes Scenic Drive 6.5 mi NNW of ranger station, west of old tram, hardwood hammock, slough and strand, March 2009, (F, holotype). Etymology.The epithet refers to the pseudostromatic clusters of aggregate lirellae. Distribution and ecology.This species is known from a single collection from Fakahatchee Strand Preserve State Park, found on branches of hardwood. to represent another species with pseudostromatic clusters of lirellae (see above). However, has muriform ascospores and its thallus is ligher, with a less conspicuous contrast between thallus and pseudostromatic clusters. appears to be the only species known in the genus with pseudostromatic clusters and transversely septate ascospores. Common & Lcking, new species Figure 26B Mycobank #560007 Diagnosis.Sicut sed ascosporis maioribus et paraphysoidis spinulosis mm longae, 0.25 mm latae. Ascosporae muriformes, 15 6 m, I. Acidi lichenum desunt.

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146 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Description.Thallus corticolous, 1 cm diam., 30 m thick, continuous; surface smooth, yellowish-brown to olive-brown. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and scattered clusters of crystals. Lirellae straight to curved, unbranched to sparsely complete thalline margin, 1 mm long, 0.3.5 mm wide, 0.15.2 mm high; disc concealed; labia conspicuous but not thickened, grayish black to brown-black below corticate thalline margin; thalline margin olive-brown. Excipulum entire, brown-black, 25 m wide; covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 80 m high, colorless, clear; epithecium granulose, 5 m high, gray-brown. Paraphyses unbranched, apically spinulose; periphysoids present, apically spinulose; asci fusiform, 80 20 m. Ascospores 2 per ascus, ellipsoid, transversely 3-septate, 25 11 m (including 1.5 m thick wall), 2 times as long as wide, with trypethelioid septa and lumina, colorless, I+ violet-blue. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (MSC, holotype). Etymology.We honor Edward Tuckerman for his contributions to North American lichenology with this new species. Distribution and Ecology.This species is known from two rich collections on hardwood from Fakahatchee Strand Preserve State Park. Remarks. externally resembles (Nyl.) Nyl. and Staiger (Staiger 2002) in the corticate, smooth, oliveAnatomically, however, the species is quite unique in having unusually large, albeit 3-septate ascospores in numbers of 2 per ascus and apically spinulose paraphyses and periphysoids. Spinulose paraphyses do occur in a few other species in (Staiger 2002), but not in combination with carbonized lirellae (except for the new species (formally described in Lumbsch et al. 2011). The ascospores are the largest 3-septate ascospores thus far known in Staiger (2002) gives (Nyl.) Staiger as species similar to F. but with larger ascospores; however, clearly differs in the largely endoperidermal, pale yellowish-gray thallus and the only weakly and apically carbonized lirellae; it also lacks spinulose paraphyses and periphysoids and is thus is quite distinct from Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (hb. Common). Common & Lcking, new species Figure 26C Mycobank #560008 Diagnosis.Sicut sed ascospores 7-septatis differt. Thallus pro parte 0.1 mm latae. Ascosporae 5-septate, 12 4 m, I. Acidi lichenum desunt. Description.Thallus corticolous, 1 cm diam., 40 m thick, continuous; surface smooth to uneven, yellowish white. Photobiont trentepohlioid. Thallus in section with irregular and loose upper cortex, irregular algal layer and clusters of crystals. Lirellae straight to curved, unbranched to sparsely branched, immersed, long, 0.1 mm wide, 0.1 mm high; disc concealed; labia inconspicuous; thalline margin yellowish white but yellow-brown along the slit. Excipulum entire, colorless to apically orange-brown, 15 m wide; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 5 m high, colorless to pale yellowish; hymenium 60 m high, colorless, clear; epithecium granulose, 5 m

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 147 high, gray-brown. Paraphyses unbranched, glabrous; periphysoids not observed; asci fusiform, 60 10 m. Ascospores 8 per ascus, ellipsoid, transversely (3-)5-septate, 12 4 m (including 0.5 m thick wall), 3 times as long as wide, with trypethelioid septa and lumina, colorless, I. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Big Cypress Bend, old growth Cypress swamp, Apr 1997, (MSC, holotype). Etymology.The name refers to the transversely multiseptate ascospores. Distribution and ecology.This species is known from a single collection, growing on hardwood, from Fakahatchee Strand Preserve State Park. Remarks. is and perhaps closely related to that species. It differs, however, in the mostly 5-septate ascospores, a highly unusual trait in the genus, where the species known to date have either 3-septate or (sub-) muriform ascospores (Staiger 2002). In spite of careful search, we have not found any indication in the literature or studied type specimens that suggest a previous description of a multiseptate species that would belong in On the other hand, most species with muriform ascospores have 5 transverse primary septa. Common & Lcking, new species Figure 27DE Mycobank #560009 Diagnosis.Sicut sed ascosporis minoribus et thallo albo-cinereo differt. Description.Thallus corticolous, 1 cm diam., 50 m thick, continuous; surface smooth, white to pale gray. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and large clusters but often in radiating lines, immersed to erumpent, with lateral to apically thin complete thalline margin, 3 mm long, 0.1.2 mm wide (abruptly thinner at the ends), 0.15.2 mm high; disc concealed; (when covered by thin thallin margin); thalline margin white to pale gray. Excipulum crenulate, apically carbonized and laterally and basally orange-brown, 30 m wide and basally to 25 m high; laterally covered by corticate algiferous thallus including clusters of crystals; hypothecium prosoplectenchymatous, 10 m high, colorless to pale yellowish; hymenium 100 m high, colorless, clear; epithecium granulose, 5 m high, gray-brown. Paraphyses unbranched; asci fusiform, 80 30 m. Ascospores 8 per ascus, oblong, transversely 9-septate, 40 9 m, 4 times as long as wide, colorless, I+ violet-blue. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The name refers to the peculiar shape of the lirellae, which become abruptly narrow and with entire labia towards the ends and hence appear appendiculate. Distribution and Ecology.Surprisingly this new species is one of the most commonly collected graphids at Fakahatchee but not yet known from outside the area, making it a possible local endemic. It mostly grows on small branches of hardwoods in partially exposed microhabitats. Remarks.With a number of over 300 accepted taxa and well over 1000 described names, describing new species in the genus is problematic. Nevertheless, in spite of careful search, we have not found a name for this taxon, which is represented by abundant material. With the immersed-erumpent lirellae with striate labia and apically carbonized excipulum, together with the clear hymenium, transversely septate ascospores, and no chemistry, it comes closest to Lcking, Redinger, Mll. Arg., and (Lcking et al. 2009).

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148 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) However, both and have ascospores over 80 m long, whereas in G. and they are 20 m long. The often present thin cortex covering the labia apically is also found in but besides the larger ascospores, that species also differs by its dark, olive-gray thallus. can be considered a further species complex. is the most abundant at Fakahatchee. The collections made during the Tuckerman workshop in March 2009 are almost without exception sterile, whereas those made by R. Common in Apr 1997 are abundantly fertile, suggesting not just a seasonality in ascospore production but periodicity that could span more than a years cycle. The species does not key out in Harris (1995); the most similar species there is Nyl., which has prominent to sessile lirellae and smaller ascospores, but it too is listed by Harris (1995) as frequently sterile. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (hb. Common); ibid., K2 trail among royal palms, second growth area, Apr 1997, (all hb. Common). U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, off James Scenic Drive 6.5 mi NNW of ranger station at gate 12 along old tram road, hardwood hammock, strand and swamp, on bark, March 2009, (hb. Lay), , (all F, USF). Common & Lcking, new species Figure 44D Mycobank #560010 Diagnosis.Sicut sed acidum sticticum continente differt. Description.Thallus corticolous, 1 cm diam., 50 m thick, continuous; surface uneven-verrucose, pale yellow-gray to oliveyellow. Photobiont trentepohlioid. Thallus in section with prosoplectenchymatous upper cortex, irregular algal layer and large clusters of crystals. not pseudostromatic, stellately branched, erumpent, with indistinct labia and thin thalline margin, 1 mm long (but sometimes secondarily divided into shorter segments), 0.1 mm wide, 0.12.15 mm high; disc exposed, chocolate-brown, sometimes with thin gray pruina; labia indistinct, dark brown; thalline margin yellow-white. Excipulum divergent, dark brown, 10 m wide; laterally covered by corticate algiferous thallus with clusters of crystals between individual lirellae; hypothecium prosoplectenchymatous, 10 m high, colorless to pale yellowish; hymenium 50 m high, colorless, clear; epithecium granulose, 5 m high, gray-brown. Paraphyses unbranched; asci fusiform to clavate, 45 10 m. Ascospores 8 per ascus, oblong, transversely 3-septate, 13 5 m, 2 times as long as wide, colorless, I+ vine-red to red-purple. Secondary chemistry: stictic acid (major), constictic acid (trace), thallus section Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (MSC, holotype; hb. Common, isotype). Etymology.The name refers to the delicate lirellae. Distribution and Ecology.Known from three collections from Fakahatchee Strand Preserve State Park, all on branches of hardwood. Remarks.The clear hymenium, aggregatestella te lirellae, and small ascospores indicate this new species to belong in the aggregate (Staiger 2002). Most species in this aggregate have norstictic acid; thus far only one species, is known with stictic acid (Dal Forno & Eliasaro 2010). The latter has more robust, distinctly pruinose lirellae and larger, usually 5-septate ascospores. Additional Specimens Examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, K2 trail among royal palms, second growth area, Apr 1997, (hb. Common), (hb. Common).

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 149 Common & Lcking, new species Figure 56C Mycobank #560011 Diagnosis.Sicut sed apotheciis marginibus pruinosis et ascosporis minoribus differt. Description.Thallus corticolous, 1 cm diam., 30 m thick, continuous; surface uneven, white to pale gray. Photobiont chlorococcoid. Apothecia sessile, rounded, 0.3.7 mm diam. and plane to slightly convex in old apothecia, black; margin thick, prominent, evanescent in old apothecia, thickly gray-pruinose especially in younger apothecia. Excipulum paraplectenchymatous, 30 50 m broad, dark purplish-brown. Hypothecium 30 m high, dark purplish brown, K+ purplish. Apothecial base dark purplish brown. Epithecium 10 m high, blackish brown. Hymenium 100 120 m high, colorless or light purplish in upper parts. Paraphyses branched and anastomosing. Asci 9010 18 m. Ascospores 4 per ascus, ellipsoid, muriform with 3 transverse and 0 longitudinal septa per segment, 20 9 m, 2.5 times as long as broad, colorless. Campylidia sessile, 0.4.5 mm broad, 0.6.8 mm long; lobe well-developed, hood-shaped, 7-septate, 80 2.5 m. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (MSC, holotype, USF, hb. Common, isotypes). Etymology.Refers to the state of Florida. Distribution and Ecology.This species is known from two, well-developed collections from Fakahatchee Strand Preserve State Park, growing on branches and trunks of hardwood. Remarks. genus characterized by lecideine apothecia, -type asci, anastomosing, net-like paraphyses, thin-walled, transversely septate to muriform ascospores, and campylidioid conidia (Lcking 2008). The species are and size and the absence or presence of pruina on the apothecial margin. Thus far, only one species, was known to produce small, muriform ascospores in numbers of 4 per ascus (Lcking 2008). species agrees with in ascospore type, but the young apothecia are epruinose in and thickly gray-pruinose in the new species; in addition, the ascospores are larger in (25 14 m). Additional Specimen examined. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Scenic Drive (CR 837) just past bend near gate 14, second growth, Apr 1997, (hb. Common). Lcking & Rivas Plata, new species Figure 56D Mycobank #560012 Diagnosis.Sicut sed thallo granuloso differt. Description.Thallus corticolous, 2 cm diam., 30 m thick, continuous; surface granules (isidia) formed by agglomerated algal cells wrapped in fungal hyphae. Photobiont chlorococcoid. Apothecia sessile, rounded to irregular in outline, 0.5 mm diam. and 180 270 m high; disc plane becoming convex, black to brownish black; margin thin but distinct, not prominent, persistent, black. Excipulum paraplectenchymatous, 30 m broad, graybrown. Hypothecium 50 m high, dark purplish brown, K+ purplish. Apothecial base dark purplish brown. Epithecium 10 m high, blackish brown. Hymenium 100 m high, colorless or light purplish in upper parts. Paraphyses branched and anastomosing. Asci 90 18 m. Ascospores 4 per ascus, ellipsoid, muriform with 3 transverse and 1 longitudinal septa per segment, 20 10 m, 1.8 times as long as broad, colorless. Campylidia sessile, 0.4.6

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150 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) mm broad, 0.7 mm long; lobe well-developed, hood-shaped, dark gray to black with white-gray 5-septate, 40 2 m. Secondary chemistry: no substances detected by TLC. Type. U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Janes Scenic Drive 1.8 mi NNW of ranger station, hardwood prairie hammock, March 2009, (F, holotype; USF, isotype). and densely granulose thallus, an unusual feature in the genus Distribution and Ecology.This species is known from two well-developed collections from Fakahatchee Strand Preserve State Park, growing on bark. Remarks.This is a second new species related to which is also found at the type locality. Like the preceeding species, shares with the small, muriform, broadly ellipsoid ascospores occurring in numbers of 4 per ascus. However, differs from all species known to date. It is unlikely that these granules serve as isidia; rather they should be interpreted as increasing the thallus surface structure is known from the related genus Lasioloma (Lcking & Srusiaux 2001). Additional Specimen Examined.U.S.A. Florida. Collier Co.: Fakahatchee Strand Preserve State Park, Janes Scenic Drive 1.8 mi NNW of ranger station, hardwood prairie hammock, March 2009, (hb. Lay), (F), (UPR). ANNOTATED CHECKLIST OF LICHEN SPECIES FROM FAKAHATCHEE STRAND PRESERVE STATE PARK In the following list, genera and species are listed in alphabetical order. Brief taxonomic and nomenclatural annotations are given for selected taxa. Selected and representative specimens are cited for each taxon in parentheses after the species name. Herbaria are not indicated in the list but are as follows: DUKE (B. P. Hodkinson), F (R. Lcking & E. Rivas Plata, M. P. Nelsen), FTU (W. Safranek), ILLS (L. Crane), LI (O. Breuss), MSC (R. Common, types and new records for United States), NY (S. Q. Beeching p.p., W. R. Buck, R. C. Harris, J. C. Lendemer), OAC (T. McMullin), UPR (J. A. Mercado-Daz), and the private herbaria of S. Q. Beeching, R. Common, M. Hodges, E. Lay, W. B. Sanders, H. P. Schaefer, and R. & J. Seavey. Where the material allowed, selected duplicate specimens are deposited at the University of Southern Florida in Tampa (USF); these are marked with an asterisk (*) after the collection number. Most crustose species are illustrated by color photographs in a series of plates in the supplemental online appendix (Figs 4; http:// www.flmnh.ufl.edu/bulletin/vol49no4supplmats. htm); for a few taxa for which the Fakahatchee material did not allow to produce good images, we replaced these by images taken from specimens from other regions. For selected genera that are either not treated in detail in Harris (1990, 1995) or more recent publications (Ekman 1996; Staiger & Kalb 1999; Marbach 2000; Lcking et al. 2007; Lendemer et al. 2009; Seavey 2009; Seavey & Seavey 2009; Tripp et al. 2010) or for which many new records are cited, we provide brief keytables that summarize the characters of each species and these keytables, species are arranged in taxonomic order by column characters from left to right; to key out a species, one would therefore start with the North American lichen checklist are marked with an asterisk (*) before the species name; some of these have already been known to occur in North America but have either not been reported in the literature or not yet added to the checklist. Species newly reported to the New World are proceded by two asterisks (**) and new species (described above or in separate papers) with three asterisks(***) (M. Wirth & Hale) E. A. Tripp & Lendemer Figure 4AB

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 151 ( , , *, *, *). Material from Florida was sequenced and clusters with aff. from Cuba (Staiger et al. 2006; Rivas Plata et al., unpubl. data). Thus far, no other species of have been sequenced, but based on the available data, the genus is distinct from other lirellate Graphidaceae and unrelated to and instead close to (Nyl. Tuck.) E. A. Tripp & Lendemer Figure 4C ( ). Species of are recognized by their non-carbonized lirellae, warty paraphysis tips and usually non-amyloid ascospores. A taxonomic treatment of North American species is given by Tripp et al. (2010). spec. Figure 4D ( ). This material represents an undescribed foliicolous species most probably belonging in (Lcking 2008). It produces abundant, very shortly stalked hyphophores that carry a rounded mass of agglutinated diahyphae interspersed with algal cells. No apothecia were found in the material. (Malme) Marbach Figure 4E ( *). Species of are treated in Marbach (2000). The genus is probably heterogeneous and needs a revision using molecular methods. (Hoffm.) Coppins & Scheid. Figure 4F ( *). (Zahlbr.) R. C. Harris ( ). R. C. Harris ( ). Harris (1995) provided a key to species, both for Florida and world-wide. R. C. Harris ( ). (Nyl.) R. C. Harris Figure 4G ( ). (Nyl.) R. C. Harris igure 4H ( ). (Nyl.) R. C. Harris ( ). spec. Figure 5A ( *, ). The material agrees with in Harris (1995) and probably represents an undescribed species. Unfortunately the material is not abundant enough for a formal description. (Zahlbr.) R. C. Harris ( ). Species of are treated in Harris (1995) (Eschw.) R. C. Harris Figure 5BD ( , *, *, , ). (Fe) Nyl. Figure 5EF ( ). The genus is not well-understood and the present study revealed several additions to the North American lichen checklist; some of these are widespread in the coastal plain but have not yet been reported in the literature. The taxonomy of the species found here is summarized in Table 2. Compare also (DC.) Wallr. Figure 5GH ( , *). As pointed out by with variation in ascomata morphology and ascospore size and septation. In our material, has few marginal pruina and the ascospores are mostly 4-septate and 20 6 m. Fe Figure 6A ( ). Taylor Figure 6B ( description given in Grube (2007), particularly with remnants of bark tissue resembling a

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152 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) cited in this work (only lichenized species included). Septa = number of ascospore septa: muri = muriform; Shape = ascospore shape: iso = isolocular (cells equal or middle cell sometimes slighly enlarged), macro = macrocephalic (upper and sometimes also lower end cell enlarged), submacro = submacrocephalic (second upper cell enlarged); Ascomata = shape of ascomata: irreg = irregular; Color = color of ascomata; Pruina = pruinose cover of ascomata; Emergence = emergence of ascomata; Chemistry = secondary thallus chemistry (not including ascomata pigments): lichexan = lichexanthone, psor = psoromic acid; Other = other characters: inspersed = inspersed hymenium, K+ pigments = crystallized pigments present in excipulum, epithecium and/or hymenium, segmented = lirellae divided into segments; Size = ascospore size in m (typical upper range for length and size; the typical lower range is about 30% less). Detailed hymenial and ascus reactions with iodine solution are not given for lack of space (consulting specialized literature for species is highly recommended; e.g. Grube 2007). Species of Septa Shape Ascomata Color Pruina Emergence Chemistry Other Size 3 submacro irreg-lirellate pale adnate lichexan 18 5 A. aff. 3 submacro rounded-irreg pale pruinose erumpent 28 10 3 macro irreg-lirellate cinnabar pruinose erumpent K+ pigment 18 6 3 macro irreg-lirellate cinnabar pruinose erumpent psor K+ pigment 22 6 4 macro irreg cinnabar pruinose erumpent K+ pigment 28 8 4 macro lirellate brown-orange erumpent K+ pigment 20 8 3 submacro irreg dark red adnate K+ pigment 25 12 3 macro lirellate brown adnate segmented 18 6 4 macro lirellate-stellate brown adnate 30 12 3 submacro irreg black pruinose adnate 15 5 5 submacro irreg-lirellate black adnate 15 6 5 macro rounded-irreg black adnate 25 8 5 macro rounded-irreg brown-black bark tissue erumpent 35 12 11 iso rounded-irreg black adnate 70 20 11 iso rounded-irreg black adnate inspersed 60 15 muri iso rounded-irreg black adnate 30 10 muri iso rounded-irreg black adnate inspersed 70 30 muri iso rounded-irreg black adnate K+ pigment 50 20 muri iso irreg-lirellate brown-black adnate K+ pigment 55 20

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 153 coarse pruina and later becoming erumpent to almost sessile and -like. ** Nyl. Figure 6CD ( ). Fe Figure 6EF ( , *). Nyl. ( ). The description for the North American material of this species in Lendemer et al. (2009b) is reminescent of another, recently described species, Grube, which was also main difference is in ascospore size (less than 50 m long in versus more than 50 m long in ) and the often more lirellate ascomata in ; however, it case would be the older name. Grube Figure 6GH ( ). (Tuck.) R. C. Harris ( ). This name is here applied to a species that apparently belongs in the genus (and therefore not included in Table 2), with 8-spored asci and 2O acids. This taxon has been confused with the recently described (Seavey 2009), which also has 8-spored asci but smaller ascospores and contains psoromic acid and lichexanthone. The name Tuck. (Tuckerman 1872) is apparently based on material that Fe, based on an Auber collection from Cuba. We have not yet seen that material and in the light of the chemical diversity of species, it is possible the taxon from southeastern North America named here in which case a new species name would be required. For this reason, we have refrained at this time from simply transferring to aff. Anzi Figure 7A ( name for this taxon which has macrocephalic, 3-septate ascospores about 15 5 m in size and abundant pycnidia and appears to be related to the bulk of foliicolous taxa (Lcking 2008). in most details, including the abundant pycnidia (Grube 2007), except that it is lichenized. Nyl. Figure 7B ( *, *, ). aff. (Pers.) Steud. A. L. Sm. Figure 7CD ( ). This material deviates from typical (Grube 2007) by its larger ascospores (24 m). Nyl. Figure 7EF ( ). (Fe) Nyl. Figure 7GH ( ). Grube Figure 8AB ( ). (Fe) Nyl. Figure 8C ( *). G. Merr. Grube & Lendemer Figure 8D ( , *, ). This name was recently validated (Grube & Lendemer 2009). ** Nyl. Figure 8EF ( ). We apply this name as it well in ascomata morphology and anatomy and ascospore type. However, it is likely that there are additional names available for this relatively non-descript taxon. (Mll. Arg.) Grube Figure 8GH ( ). spec. ( ). This is an unnamed, yet characteristic taxon. The ascomata are black, rather plane, and irregular in shape; the hamathecium is slightly inspersed with oil

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154 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) droplets and reacts K (no crystalline pigments present); ascospores are muriform, about 35 40 m in size and become brown when mature. is most similar but has much larger, hyaline ascospores. The African Nyl. also resembles this material but its ascospores are persistently hyaline and have an enlarged distal cell. (Ach.) Mll. Arg. Figure 9A ( ). Species of are keyed out in Harris (1975, 1995). R. C. Harris Figure 9B ( ). (Ach.) Mll. Arg. Figure 9C ( ). aff. (Ach.) Mll. Arg. Figure 9D ( ). This material agrees with in ascospore type but has consistently excentric (lateral) ostioles. We have not found a name for this taxon but refrain from describing it as new, since the genus is in further need of revision and possible epithets might be hidden among other generic names. A. Massal. Figure 9E ( *, *, ). This species strongly resembles an with muriform ascospores, but differs in the strongly compacted, stromatic structure of the hymenium in which the asci are formed in locules and surrounded by a delicate network of anastomosing paraphysoids. The entire hymenium in the present material appears inspersed. spec. ( the described species in the genus. Vain. Figure 9F ( ). For key to species of see Lcking (2008). (Malme) R. Sant. Figure 9G ( ). Lcking Figure 9H ( *). Mll. Arg. Figure 10A ( ). This genus is treated in Harris (1995). Molecular data suggest that the genus (1995) idea of including the bulk of species of , , and within a single genus (or ) is correct (Nelsen et al. 2009). In addition, the species concept in the / aggregate needs revision. We do not agree with Harris (1995) that species with either fused or separate ostioles should be lumped, as there are substantial differences in other morphological features as well. In addition, hymenial inspersion has not yet been explored as a taxonomic character in this complex, even if its use is accepted for other genera. Mll. Arg. Figure 10B ( *, *). This species is quite distinctive due to its very large ascospores (70 17 m) which, however, have only 5 septa with distinctly diamond-shaped lumina, thus looking like gigantic ascospores of the -type. The thallus and especially the perithecial warts, which contain 3 aggregate perithecia with fused ostioles, are UV+ golden-yellow (lichexanthone). The species was described from Cuba (Mller Argoviensis 1885), so its presence in Florida is not a surprise. Nyl. Figure 10C ( , , ). Kremp. Figure 10D ( , ). Mll. Arg. Figure 10EF ( *, *, *, ). This taxon is supposed to have a clear hymenium, but Harris (1995) also included specimens with inspersed hymenia. We found one collection with an inspersed hymenium

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 155 ( ) and suspect that this represents a separate taxon, but no name appears to be available for this form. (Vain.) R. Sant. Figure 10G ( ). Lcking (2008) provided a key to species of (Stirt.) R. Sant. Figure 10H ( ). Malme. Figure 11A ( *). Species of (Tuck.) S. Ekman & Kalb Figure 11B ( *, *, ). (Mll. Arg.) Zahlbr. Figure 11C ( *, ). Bacidia aff. (Mll. Arg.) Zahlbr. Figure 11D ( ). This material differs from gray margin. (Nyl.) Zahlbr. Figure 11E ( , *, ). Bacidia aff. (Nyl.) Zahlbr. Figure 11F ( ). This material keys to in Ekman (1996) but differs in the dark apothecia without orange tinge and the persistent margin. (Tuck.) Zahlbr. Figure 11G ( *, *, ). Bacidia aff. (Tuck.) Zahlbr. Figure 11H ( ). This material resembles but has darker apothecia and the internal parts (excipulum) have an orange, K+ purple band. The material is also similar to in which the excipulum is pale orange and not K+ purple. Malme. Figure 12A ( *, ). (Kremp.) Zahlbr. Figure 12B ( *, ). (Fr. E. Michener) A. Schneid. Figure 12C ( *). Bacidia aff. (Fr. E. Michener) A. Schneid. Figure 12D ( ). This material covers the taxon with brown apothecia and brown epithecium included by Ekman (1996) in In our view, this is a taxon separate from s.str. with black apothecia and red-black hypothecium, but there appears to be no alternative name available. The type material of all synonyms cited under B by Ekman (1996) seem to represent the typical form with black apothecia. We have refrained from formally describing the taxon with brown apothecia since our material is moderatly developed and segregating this taxon requires restudy of the many specimens cited by Ekman (1996). Bacidia spec. 1 Figure 12E ( ). This material does not key to any species in Ekman (1996). The apothecia are large, brown and irregular in outline, with an orange excipulum and hypothecium. The ascospores are 40 4 m and thus unusually broad, with 51 septa. Most similar in ascospore type are Zahlbr. and B. (Nyl.) Arnold, which have black apothecia with different apothecial pigments internally (Ekman 1996). Bacidia spec. 2 ( ). The taxonomic status of this material is unclear. The granular, green thallus and the more or less emarginate, turbinate to convex, dark gray-brown apothecia, together with the 3-septate ascospores, suggest a species of or but none of these two genera includes anything similar. (De Not.) Coppins agrees in general morphology but has larger ascospores. Since the ascospores of the present material are rather narrow (to 13 2.5 m) and the excipulum consists of parallel hyphae, is a likely home, but no matching species was found in the key provided by Ekman (1996). S. Ekman Figure 12F ( ).

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156 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) (Vain.) Egea & Torrente Figure 12G ( ). The two species of found at Fakahatchee are characterized by cylindrical ascospores lacking constrictions and a K/I, rather thick excipulum (Egea and Torrente 1993). has a denticulate apothecial margin and ascospores to 130 m long, whereas has a smooth margin and ascospores less than 100 m long. (Fe) Egea & Torrente Figure 12H ( , ). (Tuck.) Marbach Figure 13A ( ). Marbach (2000) provided a key to species. (R. C. Harris) Marbach Figure 13B ( *). (Tuck.) R. C. Harris Figure 13C ( ). Species of are treated in Harris (1995). (Eschw.) Trevis. Figure 13D ( ). (Spreng.) R. Sant. & Hafellner. Figure 13E ( *, ). J. Steiner ( ). (Vain.) Zahlbr. Figure 13F ( ). Lcking (2008) provided a key to neotropical species including those found in Florida. (Nyl.) Vain. Figure 13G ( *, ). (Tuck.) S. Ekman Figure 13H ( , , ). (Nyl.) P. James ( ). Nyl. Figure 14AB ( ). Figure 14C ( ). This recently described species (Lumbsch et al. 2011) is closely related to but differs in the heavily pruinose apothecia. In contrast to the apothecia disc is orange-brown and the pruina yellowish, and also the apothecial margins are pruinose. *** Calopadia Hodges & Lcking Figure 14DG ( ). See p. 138 for description of this new species. The species of Calopadia currently known from Florida are summarized in Table 3. Figure 14H ( , *). *** Calopadia imshaugii Common & Lcking Figure 15AF ( ). See p. 139 for description of this new species. Figure 15GH ( , ). Figure 16A ( ). Calopadia cited in this work. Ascospore length in m. Species of Calopadia Disc color Pruina Hypothecium color Ascospore number Ascospore length C. imshaugii orange-brown white-gray brown (1)2()/ascus 50 orange-brown yellowish light brown single 8010 orange-brown yellowish light brown single 50 gray-brown white-gray dark brown single 70 orange-brown light brown single 60 gray-brown dark brown single 55 black dark brown single 60 black aeruginous single 55

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 157 Figure 16B ( , , *, , ). Figure 16C ( ). Calopadia spec. ( ). This material bears only campylidia and cannot be to its conspicuously verrucose thallus, a feature uncommon in Calopadia and the related genus The campilidioconidia associate with photobiont cells, a feature more commonly observed in the genus (Lcking 2008). Caloplaca camptidia (Tuck.) Zahlbr. Figure 16D ( ). This is one of the brown species, characterized by the pruinose apothecia resembling those of a (Wetmore 1994). (Taylor) C. W. Dodge Figure 16E ( , , *). This species is characterized by the orange and densely isidate thallus. (Hoffm.) A. E Wade Figure 16F ( ). This material features mostly deformed apothecia with brownish color, but some present corresponding to those typically found in this species. Hale ( , ). ( ). (Fe) Kalb Figure 16G ( *). This material was since the ascospores appeared to lack septa. However, some 1-septate ascospores were eventually found, and the apothecial anatomy, with the outer excipulum consisting of radiating cell rows, the inner excipulum being of medullary structure, and the hypothecium being pale, as well as the aeruginous epithecium, characterize the material as (Kalb 2007). Species of can be remarkably similar in appearance, especially as and related taxa also have a medullary excipulum, but apart from the non-septate ascospores, these species differ in the outer paraplectenchymatous excipulum and the brown-black hypothecium. and have been synonymized with (Fryday & Lendemer 2010). We do agree that the limits between the three genera are not clearcut but differences between the type species, including in ascus structure, exist. Unfortunately, no sequence data are available for ; analysis of the available sequences of the large subunit of the nuclear ribosomal DNA (nuLSU) of one specimen of and two specimens of (Lcking, unpubl. data) show that both are in the same clade within Ramalinaceae, but on very long branches each, which supports a close relationship but not congenerity, considering that nuLSU is a relatively conserved gene partition and hence species within the same genus are expected to appear on short branches. Therefore, we maintain the genera separate pending further studies. The overlap of anatomical and chemical features between species assigned to the different genera documented by Fryday & Lendemer (2010) is not necessarily evidence to merge them, since phylogenetic studies show that genetically distinctive clades can evolve similar phenotypic characters in parallel (Rivas Plata et al. 2011). (Nyl.) Vain. Figure 16H ( ). (Ach.) Mll. Arg. Figure 17A ( , , *). R. C. Harris ( , *). (Nyl.) Rivas Plata &

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158 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Mangold Figure 17B ( ). The material agrees perfectly with the type of this species in the morphology of the thallus and apothecia, which are somewhat intermediate between Chapsa and as well as the brown excipulum and hardly discernable periphysoids. The ascospores are, however, larger than in the type and material from Costa Rica (25 m versus 15 m) and with more numerous septa (9 versus 5). Since only few specimens are known, we refrain from considering the Florida material a separate taxon, until the variational amplitude of the Florida populations has been established in comparison with the material from Costa Rica and Panama (Hale 1978). (Tuck.) A. Frisch Figure 17C ( , *, *, *). This species is commonly found on the thin trunks and branches of swamp dogwood. Chapsa (Tuck.) Breuss & Lcking Figure 17D ( *, *). See p. 140 for new combination and discussion. ** (Hale) Mangold ( Lanka well in the endoperidermal, pale thallus, the irregular apothecia, the amyloid ascospores with rather thin septa, and the lack of secondary substances. Previously known from Sri Lanka and Australia (Hale 1981; Mangold et al. 2009); new for the New World. (Vain.) Kalb Figure 17E ( , , ). (Tuck.) Fink ( ). (Flrke Sommerf.) Spreng. ( , *, ). (Fe) Vain. ( , ). var. (Zoll. & Moritz) Vain. ( ). Tuck. ( ). (Vain.) Sandst. ( , , *, *). R. C. Harris Figure 17F ( , ). comprises species of s.lat. with thallus-dominated perithecial verrucae and shiny thallus and (dark) prothallus; the Florida species are treated in Harris (1995). (Malme) R. C. Harris Figure 17G ( , *). (Ach.) R. C. Harris Figure 17H ( ). Vain. Figure 18A ( ). (Spreng.) Swinscow & Krog Figure 18B ( , ). (Spreng.) Arv. & D. J. Galloway Figure 18CD ( , , *). C. W. Dodge Figure 18E ( ). The genus (including ) is treated with a worldwide key including all Florida species by Rivas Plata et al. (2006). (P. Henn) Lcking Figure 18F ( ). Nyl. Figure 18G ( ). Ehrenb. Figure 18H ( ). Rivas Plata, Lcking & Umaa. Figure 19A ( ). (Dicks.) Kalb & Lcking

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 159 Figure 19B ( , ). Rivas Plata, Lcking, Umana & Chaves. Figure 19C ( ). Farkas) Lcking, Aptroot & Sipman. Figure 19D ( *). (Arnold) Du Rietz ( ). Ach. ( ). (Mll. Arg.) Grube Figure 19E ( ). (Fe) Marbach Figure 19F ( ). (Sw.) A. Massal. Figure 19G ( *). Nyl. Figure 19H ( , *, ). (Tuck.) D. Hawksw. & Dibben Figure 20A ( ). (Mll. Arg.) R. Sant. Figure 20B ( ). The otherwise well-developed material has mostly sterile ascigerous parts but in two specimens a few young asci where found with eight undeveloped ascospores per ascus. Seavey Figure 20CD ( ). *** Vain. Lcking Figure 20EF ( *). See p. 140 for description of this new species. Sparrius. Figure 20G ( , ). In this material and other specimens from the Neotropics, the entire thallus contains gyrophoric acid and reacts C+ red, not just the soralia as in the paleotropical populations (Sparrius & Saipunkaew 2005). G. Thor Figure 20H ( , , *, , ). This species is often misinterpreted in the literature as having isidia. Isidiate with a chemistry similar to that of C. (gyrophoric acid) must be referred to (Vain.) Aptroot & Lcking (Aptroot et al. 2009). is one of the most common and widespread species of the genus in the Neotropics and consistently fertile, always lacking isidia. (Mll. Arg.) Zahlbr. Figure 21A ( ). f. phyllophilum Parm. Figure 21B ( ). (Mont. & Bosch) Kalb, Figure 21C ( , ). This genus has been monographed by Kalb et al. (2004) and North American species are treated in Tripp et al. (2010). *** M. Cceres & Lcking Figure 21D ( ). This species, described in a separate paper (Lumbsch et al. 2011), was (Nyl.) Staiger (Cceres 2007), with which it agrees in the small ascospores and norstictic acid but differs in the absence of hymenial inspersion and the indistinctly corticate thallus. Sequencing data showed that the species belongs in instead, with which it agrees in the thallus and ascomata morphology (Lumbsch et al. 2011). Figure 21E ( *). (Afz.) B. J. Moore Figure 21F ( , *). D. D. Awasthi Figure 21G ( ). (Stein) D. D. Awasthi Figure 21H ( *). (Nyl.) D. D. Awasthi Figure 22AB ( ). (Sw.) Schaer. Clem. Figure

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160 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) 22C ( , ). (Vain.) B. Moore Figure 22D ( *). (Ach.) A. Massal. Figure 22E F ( , , , , ). Lcking & W. R. Buck. Figure 22G ( , ). cf. Fe ( ). The material is sterile but suggests this species based on the verrucose thallus, white setae and acute white hyphophores with subapically inserted diahyphal bunches. (Mll. Arg.) R. Sant. ( ). aff. (Mll. Arg.) R. Sant. Figure 22H ( ). This material represents an undescribed species, but the single thallus is too small to describe it formally. It agrees with the pantropical E. in the short hyphophores with brown-black upper part, but contrary to the latter, the diahyphal bunches include numerous small algal cells with are dispersed with the in with this mode of joint dispersal of mycobiont and photobiont (Lcking 2008). Figure 23A ( ). Figure 23B ( *). (Nyl.) Redinger Figure 23C ( *). Figure 23D ( ). *** Common & Lcking Figure 23E ( ). See p. 141 for description of this new species. sensu Staiger (2002) is a rather large genus, with over 60 species (Lcking, unpubl. data). Species concepts have so far been confusing and the delimitation of species found in Florida is summarized in Table 4. *** Common & Lcking Figure 23FG ( , ). See p. 142 for description of this new species. (Nyl.) Staiger Figure 23H ( ). (Tuck.) Staiger Figure 24A ( , ). *** Common & Lcking Figure 24B ( , ). See p. 142 for description of this new species. ** Mont. & Bosch Figure 24CD ( , ). This species closely resembles C. Knight and Mitt., but differs in the strongly amyloid ascospores. has been reported for Florida (Harris 1995), but the specimens have to be rechecked for ascospore amyloidity. Archer (2009) synonymized under whereas Galloway (2007) kept them separate but instead listed (Nyl.) Nyl. and C. Knight & Mitt. as synonyms of This suggests some confusion about species concepts and the corresponding type material. has labiate lirellae (swollen lips), clearly seen in the picture of the type (Hayward 1977), whereas in and in addition, has a carbonized excipulum and blackened lirellae. The confusion about probably stems from Staiger (2002), who used the example of to denote carbonized lirellae in her scheme of classifying lirellae types in and also correctly depicted the excipulum as apically carbonized in the lectotype of (Staiger 2002:133, gave the excipulum as non-carbonized. The type material of on the other hand, agrees well with and

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 161 cited in this work and otherwise reported from Florida. Septa = number of ascospore septa: muri = muriform; iodine = ascospore amyloidity; chemistry = secondary chemistry; inspersion = hymenium inspersion; excipulum = excipulum carbonization; lirellae = lirellae shape; other = other characters: verrucose = thallus verrucose, endoperidermal = thallus endoperidermal; size = ascospore size in m (typical upper range for length and size; the typical lower range is about 30% less). Species of Septa Iodine Chemistry Inspersion Excipulum Lirellae Other Size 7 cortex loose 15 5 3 inspersed cortex loose 15 6 3 I+ carbonized roof-like 13 5 F. humilis 3 I+ carbonized roof-like 20 8 3 I+ carbonized roof-like 35 12 3 labiate 20 10 3 labiate-gaping verrucose 18 9 3 I+ labiate-gaping verrucose 18 8 3 labiate-aggregate 15 6 3 pseudostromatic 18 6 F. radiata 3 (I+) 15 7 3 cortex loose 16 6 3 (I+) 15 7 F. tachygrapha 3 (carbonized) endoperidermal 20 9 muri ecorticate 100 25 muri I+ psoromic roof-like 30 15 muri I+ psoromic labiate-gaping verrucose 30 18 muri carbonized 22 8 muri (I+) chroodiscoid 18 8 F. aff. muri labiate-gaping verrucose 25 15 muri I+ labiate 28 12 muri pseudostromatic 35 12 muri 30 15 muri (I+) 20 10 muri (I+) labiate-gaping 20 9

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162 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) should be considered a synonym of the latter, thus adding to the lichen biota of New Zealand. The main differences between F. and are the labiate versus versus smooth to uneven thallus. (Mll. Arg.) Lendemer ( , ). This species is most certainly not a genuine on the basis of its into the Thalloloma complex but is set apart by its I ascospores. Molecular data are required to clarify the systematic position of this taxon. (Nyl.) Nyl. Figure 24E ( ). This species is close to which differs in the erumpent lirellae with distinct labia and gaping disc, resembling rather than (Staiger 2002). in contrast, has immersed lirellae with roof-like margins and concealed to slightly exposed disc. However, intermediate forms do occur and in fact this intermediate lirellae type is quite common in several species of such as North American material with this type of lirellae has also been (Nyl.) Nyl. However, this is based on misinterpretation of the type material (Staiger 2002). Nyl. is validly described in Tuckerman (1888), who cited a collection from Cuba ( ) as the only collection (based on Nylander 1886). Three collections with the name are present in Nylanders herbarium: Cuba, (H-NYL 7459!), U.S.A., (H-NYL 7460!), and Cuba, (H-NYL 7461!). was cited as type by Wirth & Hale (1978), but cannot be the type as it is not in geographical accordance with the protologue and also was collected one year after the description (Staiger 2002). as already established by Wirth & Hale (1978). is a different with similar ascospores, but olivebrown thallus and very short and broad, widely open, almost chroodiscoid lirellae with white, erect margins. This specimen is the same taxon as Fink from Puerto Rico (Fink 1927), described as having closed lirellae, but in the type clearly showing short and wide open lirellae with erect, white margins. Finally, is yet another species with lirellae of the -type, i.e. roof-shaped and distinctly carbonized. the new species (formally described in Lumbsch et al. 2011). None of the three specimens can conclusively be designated as lectotype, since they all deviate from the protologue either in morphological or label details; on the other hand, it is unlikely that there are further specimens of this series which we have not seen, since the collections were checked directly in hb. Nylander and at that time only was on loan to Bettina Staiger. Conceivably, the listing of number 68 in the protologue (based on Nylander 1886) could be an error for number 69; however, deviates strongly in its lirellae type from the description given in Tuckerman rather well. Tuckerman (1888) obviously based his description on North American material (Florida, ) and stated that he had not seen an original description of Nylanders species. In that case, the validity of the description can certainly be questioned. Presently, we apply the name in the sense of pending further information. aff. (A. W. Archer) A. W. Archer. Figure 24F ( ). This material can be considered the muriform counterpart of It agrees well with the description and types of (synonym Staiger), except that the ascospores are consistently I. Given the taxonomic importance of this character in the family, the Florida material

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 163 likely represents an undescribed taxon (the non-amyloid counterpart of ), but the material is too scanty to allow for a formal description. (Vain.) Staiger Figure 24G H ( , ). (R. C. Harris) Lendemer Figure 25A ( ). The paraphyses in this species are often thickened at the apex, a feature not observed in the newly described *** Common & Lcking Figure 25B ( , , ). See p. 143 for description of this new species. (Zahlbr.) Lcking & Rivas Plata Figure 25C ( , *, ). *** Lcking & Rivas Plata Figure 25D ( ). See p. 145 for description of this new species. Figure 25E ( , ). (Mont.) Staiger Figure 25F ( *, ). The concept of this species is not clear. We have not been able to study the type, but according to Harris (1995), the labia are striate and ascospores to 24 m long. The material from Fakahatchee includes both specimens with striate and entire labia, and the latter are usually (= Nyl. in Harris 1995), supposedly also with shorter ascospores (15 m). Mason Hales Index Cards in US indicate a syntype of in UPS as striate, but with ascospores 13 m long. Tuckerman (1888) gave the ascospores of (the striate form) as 14 m, and Staiger (2002) included Mll. Arg. as synonym of ; the type of that species has robust, partly striate labia and ascospores about 15 m long. The situation is complicated by Archers (2009) synonymization of and with also suggested by Staiger (2002), who in addition suggests F. to be the oldest name for this complex. On the other hand, Harris (1995) maintained , and (= ) separate on account of lirellae morphology and ascospore size. A further problem is the fact that, while most forms in this complex have I ascospores, there is with distinctly amyloid ascospores. As a species in this complex as follows: (1) Lirellae short, erumpent-labiate, thallus unevenverrucose, usually yellowish brown = (ascospores I) or (ascospores I+ violet-blue). Nyl. and Tuck. are synonyms of either species, but the types have to be checked for ascospore amyloidity. (2) Lirellae long, prominent-labiate, entire or becoming striate, thallus smooth to uneven, usually olive-green, ascospores I = Since there is no apparent difference in ascospore size, the distinction between entire and striate forms cannot be maintained, which is also supported by specimens like the type of G. which bears both entire and striate lirellae. Synonyms of thus include: Nyl., Nyl., Mll. Arg., and var. Vain. *** Common & Lcking Figure 25G ( ). This new species is formally described in a separate paper (Lumbsch et al. 2011). (Nyl.) Staiger Figure 25H ( , , ). The concept of this species in the literature is not accurate. Harris (1995) and Staiger (2002) described it as having a non-carbonized excipulum and non-amyloid ascospores. However, as Tuckerman (1888) correctly noted, the type shows apical

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164 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) carbonization and this is also seen in all our collections. The ascospores are persistently small, not exceeding 15 5 m and usually about 11 m long, and are weakly to strongly amyloid. The species thus agrees with (Nyl.) Nyl. in having lirellae of type II according to Staiger (2002), but the latter has larger ascospores about 13 23 m long. (Nyl.) Staiger Figure 26A ( , ). *** Common & Lcking Figure 26B ( ). See p. 145 for description of this new species. *** Common & Lcking Figure 26C ( ). See p. 146 for description of this new species. O. E. Erikss. Figure 26D ( ). This species was included in (Aptroot et al. 1997) but molecular phylogenetic analysis suggests that is represents a separate lineage (Muggia et al. 2009). (Mll. Arg.) Lcking Figure 26E ( ). This earlier name replaces (Archer) Staiger (Lcking et al. 2009b). The taxon is identical with sp. 1271 in Harris (1995:10), as already noted by Staiger (2002). antedates Mll. Arg, by one year, but was published as a nomen nudum only (Nylander 1886). Ach. Figure 26FG ( , , , , *, ). The abundant material includes both forms with stroma bearing numerous rounded to slightly elongate lirellae, as well as forms with stroma bearing a single, radiately branched lirella. The stroma are uniform within a given individual, but thalli with intermediate forms have also been found, although they are rare. Possibly two distinct taxa are involved here and DNA studies will be needed to solve the issue. (Ach.) Staiger Figure 26H ( ). In the checklist (Esslinger 2010) as (Ach.) Nyl. cf. Fe Figure 27A ( *). The material is unfortunately sterile but agrees in the thallus and lirellae morphology as well as the clear hymenium with which has muriform ascospores. Other likely alternatives would be the less common Eschw. (ascospores transversely septate) and (Fink) Staiger (ascospores terminally muriform). The genus was keyed out with over 300 species world-wide by Lcking et al. (2009b); molecular phylogenetic analysis suggests that it actually includes two distantly related lineages (Rivas Plata et al. 2011). Eschw. Figure 27B ( , ). forms part of a misunderstood complex of species with completely carbonized excipulum and inspersed hymenium (Table 5). Specimens with small ascospores and norstictic acid represent either (disc concealed) or G. (disc exposed), whereas specimens with larger ascospores and lacking secondary substances are either G. (ascospores 30 m long) or G. (ascospores 40 m long). Mll. Arg. Figure 27C ( ). *** Common & Lcking Figure 27 DE ( , , , *, *, *, *, ). See p. 147 for description of this new species. Lcking & Umaa. Figure 27F ( *). Nyl. Figure 27G ( , ). Vain. Figure 27H (

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 165 cited in this work. Septa = number of ascospore septa: muri = muriform; excipulum = excipulum carbonization; inspersion = hymenium inspersion; labia = labia striation; chemistry = secondary chemistry; lirellae = lirellae emergence; margin = thalline margin of lirellae; other = other characters: exposed = exposed disc; green = green thallus; pruinose = pruinose labia; verrucose = verrucose thallus; size = ascospore size in m (typical upper range for length and size; the typical lower range is about 30% less). Species of Septa Excipulum Inspersion Labia Chemistry Lirellae Margin Other Size G. oshioi 5 absent entire norstictic prominent complete 30 7 7 apical entire erumpent lateral pruinose 35 8 9 apical striate erumpent comp thin 55 12 9 apical striate prominent absent green 60 12 5 lateral entire norstictic erumpent lateral pruinose 35 7 5 lateral striate lichexanth erumpent absent 30 8 71 lateral inspersed entire stictic erumpent lateral 35 7 7 lateral inspersed entire norstictic erumpent lateral exposed 40 8 7 complete entire lichexanth prominent basal 50 12 G. assimilis 71 complete entire norstictic erumpent lateral 40 8 71 complete entire norstictic erumpent comp thin pruinose 35 8 71 complete entire sessile absent 40 8 71 complete entire prominent lateral thick 40 8 5 complete entire prominent lateral verrucose 25 7 71 complete striate erumpent absent 50 12 11 complete striate erumpent lateral 65 12 5 complete inspersed entire norstictic erumpent basal 25 7 5 complete inspersed entire norstictic erumpent lateral exposed 30 8 G. 71 complete inspersed entire prominent absent 35 8 13 complete inspersed entire prominent basal 55 8 muri apical striate erumpent lateral 40 20 muri lateral entire erumpent lateral 80 25 G. muri complete striate prominent comp thin 120 20 term complete inspersed striate prominent comp thin 120 18 muri complete inspersed striate prominent comp thin 60 12 muri complete inspersed striate prominent comp thin 100 20

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166 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) , *, ). Redinger Figure 28A ( *). Lcking Figure 28B ( ). This species was already discussed in Lcking et al. (2008) for Costa Rica and is formally described in a separate paper (Lumbsch et al. 2011). Zenker. Figure 28C ( , ). Vain. LckingFigure 28D ( *). (Fe) Zahlbr. Figure 28E ( , *). We are uncertain about the correct application of the name The Florida material has very small ascospores to 25 m and the outer wall of the end cells is amyloid, a feature rarely observed in other species. The lectotype has ascospores 25 m long and size range; the end cell walls are usually not amyloid. It is possible that different species are involved here, but we want to await more detailed studies on ascospore morphology and chemistry in a wide range of species before using this character formally. Nyl. Figure 28FG ( *). Zahlbr. Figure 28H ( ). Fe ( ). Kremp. Figure 29A ( ). R. C. Harris Figure 29B ( ). M. Nakan. Figure 29C ( , , ). This in the concept of Staiger (2002), because of the absence of excipulum carbonization. However, molecular data show that species of are nested within and the differences in excipulum carbonization are merely gradual (Rivas Plata et al. 2011). is similar to the widespread tropical Fe and differs mainly in the presence of norstictic acid. Mll. Arg. Figure 29D ( , , , ). Lcking Figure 29E G ( *, *, *, ). (Mont.) Trevis. Figure 29H ( ). Leight. Figure 30A ( *, ). M. Cceres & Lcking Figure 30CF ( *, and interesting disjunction, as the species was previously only known from northeastern Brazil (Cceres 2007). The Florida material agrees well with the type, except that the ascospores are 5-septate rather than 7-septate and at the lower side of the size range (20 m versus 20 m). As in other cases, we expect this species to be also found in the Caribbean and/or parts of Central America. Lcking & Chaves. Figure 30B ( , ). Mll. Arg. Figure 30G ( *, *). (R. C. Harris) Lendemer Figure 30H ( ). Lcking & Lendemer Figure 31AB ( ). (Cavalc. & A. A. Silva) L. I. Ferraro, Lcking & Srus. Figure 31C ( ). Safranek & Lcking Figure 31DE ( ). Figure 31F

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 167 ( , ). Herrera-Campos & Lcking Figure 31G ( ). aff. W. R. Buck & Srus. Figure 31H ( ). This material resembles in the more or less radially symmetrical hyphophores with a crater-like base and the irregularly incised appendages, but differs clearly in that the diahyphae form a compact mass embedded in a gelatinous matrix. Lcking is somewhat similar but has a verrucose instead of smooth thallus and the hyphophores have a thinner margin and more delicate appendages. Unfortunately, the material is too scanty to allow for a formal description. Hillm. Figure 32A ( ). Kalb & Staiger Figure 32B ( ). (Fe) A. Massal. Figure 32C ( ). (Malme) Sheard. Figure 32DE ( ). (Malme) Marbach Figure 32F ( ). (Malme) Marbach Figure 32G ( *). *** Common & Lcking Figure 32H ( ). This new genus and species is described by Lcking et al. (2011). The genus is characterized by combining a -like whitish thallus and carbonized, labiate lirellae with a -like inspersed hymenium and gray-brown, phis -type ascospores (short with rounded ends and thick walls compared to longer with tapering ends and thinner walls in ). It is phylogenetically distinct from both and s.str., although more closely related to the latter. *** M. P. Nelsen & Lcking Figure 33AB ( , ). This new genus, with two new species, was described by Nelsen et al. (2010). It is a genus of two sterile taxa based on Aptroot et al. from the Palaeotropics (Aptroot et al. 2009). The North American species has sausage-shaped isidia and an unusual chemistry of 5-hydroxy-4-O-demethylnotatic acid and 4-O-demethylnotatic acid. (Vain.) Aptroot, Lcking & G. Thor Figure 33CD ( ). Aptroot, Lcking & Will-Wolf. Figure 33E ( *). (Ehrenb.) Aptroot, Lcking & G. Thor Figure 33FG ( , , *, ). A. Frisch & G. Thor Figure 33H ( ). This species was recently established based on material from southern Florida (Frisch et al. 2010). (Pers.) Swinscow & Krog. Figure 34A ( *, ). (Kurok.) W. L. Culb. Figure 34B ( *). (Mont.) Riddle Figure 34CD ( , , *, ). Nyl. Figure 34E ( ). Vain. Figure 34F ( *). Nyl. Figure 34G ( ). (Ach.) Malme Figure 34H ( *, *, ).

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168 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) ssp. (Nyl.) Imshaug & Brodo Figure 35A ( *). Stizenb. ( ). Lumbsch Figure 35BC ( , , *). (Tuck.) Brodo Figure 35D ( ). Fe Figure 35E ( *, *). Lumbsch Figure 35F ( ). (Spreng.) Kieffer Figure 35G ( *, *, ). spec. ( ). This material is characterized by beige apothecia with thin margin, much as in Stizenb., but has numerous small crystals in the amphithecium that dissolve in K and lacks crystals in the epithecium. The only species matching this apothecial anatomy is (Ach.) Ach., which has apothecia with much darker discs and thicker margin. (Mont. & Bosch) Staiger Figure 35H ( ). (Fink) Lendemer ( ). (Eschw.) Staiger Figure 36A ( , , ). (Rabenh.) Krb. ( , *). Nyl. ( *). (Riddle) Sierk. ( ). (Sw.) Gray ( , , , *). Vain. ( ). Sierk ( ). (Pers.) Hafellner & Bellem. Figure 36B ( , , *, , *). (Tuck.) Hafellner & Bellem. Figure 36CD ( , *, , *). (Nyl.) A. Frisch. Figure 36E ( ). The genus is treated with a key to all accepted species by Rivas Plata et al. (2010). (Nyl.) A. Frisch. Figure 36FG ( , *, ). (Eschw.) A. Frisch. Figure 36H ( ). R. C. Harris ( ). (Tuck. Nyl.) Kalb & Lcking Figure 37A ( ). Species of this genus were until recently treated under the name (Cceres 2007), but molecular phylogenetic analysis showed that the type species of is unrelated to the remaining taxa placed in that genus, and the new genus was erected for this group (Kalb et al. 2011). Most species can be keyed out using the treatment under in Cceres (2007) plus the key to Thai species of provided by Kalb et al. (2011). is currently listed as Tuck. Nyl. in the North American lichen checklist (Esslinger 2010). (Mll. Arg.) Kalb & Lcking Figure 37B ( *).

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 169 (Ach.) Kalb, Rivas Plata & Lumbsch Figure 37C ( *, , *, ). (Vain.) Kalb & Lcking Figure 37D ( *, ). (Mll. Arg.) Kalb & Lcking Figure 37E ( ). (Spreng.) Kalb, Rivas Plata & Lumbsch Figure 37F ( ). (Tuck.) Kalb, Rivas Plata & Lumbsch ( *). Kalb Figure 37G ( ). This taxon was recently described from Thailand (Kalb et al. 2011), being somewhat similar to but margin. Similar collections have been reported from Brazil (Cceres 2007) but were left unin the Neotropics apparently two species with slightly different chemistry are involved which need further study. (Eschw.) Kalb, Rivas Plata & Lumbsch Figure 37H ( *, *, ). (Tuck.) R. C. Harris Figure 38AB ( ). spec. ( ). This material is very characteristic because of its comparatively large (35 10 m), distally pointed, ornamented ascospores with markedly submedian septum. The thallus is endoperidermal and white and patchily UV+ yellow (lichexanthone). The perithecia are prominent, black, and slightly irregular although with more or less apical ostiole. It does not key out to any of the species of s.lat. in Harris (1995); because of its large, ornamented ascospores, it seems to belong in sensu Aptroot (1991), but none of the species currently accepted in that genus comes close. The recently described Aptroot has perithecia with markedly excentric ostiole and much broader ascospores with the distal end rounded (Aptroot et al. 2008). Fr. Figure 38C ( ). spec. Figure 38DG ( ). apothecia and tubular pycnidia and probably represents an undescribed species. (Pers.) Szatala. Figure 38H ( ). (Mll. Arg.) D. Hawksw. Figure 39A ( *). (Mll. Arg.) D. Hawksw. Figure 39B ( *, ). (Mll. Arg.) D. Hawksw. Figure 39CD ( *, *, ). The species resembles an especially as the ascospores are often pale to hardly pigmented at all. (at least the three species sequenced to date) has recently been shown to belong in Trypetheliaceae (Nelsen et al. 2009), which is supported by the thin, distinct, anastomosing, loosely net-like interascal hyphae. has a similar type of hamathecium and also belongs in Dothideomycetes, but in a separate clade distant from Trypetheliaceae (Nelsen et al. 2009). If ascospore pigmentation is variable in both and some species currently placed in could belong in and vice versa. The situation is complicated by similar looking species of (another clade in the Dothideomycetes) with branched and anastomosing interascal hyphae (Nelsen et al. 2009). R. C. Harris ( ). (Mll. Arg.) R. C. Harris ( ).

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170 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) (Ach.) R. C. Harris Figure 39EF ( ). Nyl. Figure 39GH ( ). R. C. Harris Figure 40A B ( , , , *). Thalli with the coarse, somewhat diffuse soralia typical of were found quite abundantly at Fakahatchee. Molecular analysis of some sorediate specimens (Rivas Plata et al., unpubl. data) showed that they are very similar to apotheciate specimens (see below). The latter agrees in most features with fertile except for the distinctly smaller ascospores and more open apothecia with irregular columella structures. is based on material from Cuba and so far only the type and an additional specimen from Panama ( ) have been found with apothecia. Coincidentally, a sorediate species with small ascospores, Kalb, was recently described from the Neotropics (Kalb 2009), which corresponds to forms with apothecia of the type and soralia of the type. Kalb (2009) indeed suggested to be the sorediate counterpart of In the present material, we have not found any specimen with apothecia and soralia on the same thallus; we therefore used for apotheciate material and for sorediate specimens, but molecular analysis of a large number of specimens could well reveal the presence of up to three taxa, s.str., and It is also possible that all sorediate North American material hitherto belongs to or even that that and are would be recognized. R. C. Harris Figure 40C ( *, ). (Nyl.) Hale Figure 40D ( , , *, ). (Tuck.) Tibell Figure 40E ( ). R. C. Harris Figure 40F ( ). (Nyl.) Mll. Arg. Figure 40G ( *). This taxon has been synonymized with (Fe) Mll. Arg., but differs in the larger apothecia with irregular-reticulate pseudocolumella; both are also genetically different. closely resembles but can be readily distinguished by the hyaline, transversely septate ascospores (brown and muriform in ). (Nyl.) Hale Figure 40H ( , *, *, *, *). The material of this taxon from as (Ach.) Hale, recently recombined as (Ach.) A. Frisch (Frisch & Kalb 2006). However, that species lacks a columella or very rarely has a rudimentary pseudocolumella, whereas the Florida material is without exception distinctly columellate. The name is available for the columellate taxon; this name is listed as synonym of in the North American lichen checklist (Esslinger 2010) but should be restored and be deleted. Vain. Figure 41A ( *). Brodo ( ). Tuck. Figure 41BC ( *, ).

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 171 Pers. Figure 41D ( *). Mll. Arg. Figure 41E ( *). R. C. Harris Figure 41F ( *, ). Mll. Arg. Figure 41G ( , ). spec. Figure 41H ( ). This material is the same as cited in Harris (1995: 7). It is characterized by robust, sessile lirellae with basally closed excipulum and slit-like disc, inspersed hymenium, and 7-septate ascospores 40 5 m in size with the two median cells slightly enlarged. Most similar appear to be Pers., with smaller, usually 5-septate ascospores and non-inspersed hymenium, and the paleotropical Ertz, with noninspersed hymenium, more delicate lirellae, and smaller, 6-septate ascospores. (Vain.) P. M. Jrg. Figure 42AB ( ). (Zahlbr.) Hale ( ). (Ach.) M. Choisy ( ). (Taylor) Hale ( , ). (Vain.) Hale ( *). (Hillm.) Hale ( *). (C. W. Dodge) Srus. ( ). (Steiner) Hale ( ). (Wulfen) A. Massal. ( ). (Nees & Flot.) Hale ( , ). (Delise Nyl.) Hale ( *). Mll. Arg. ( ). R. C. Harris Figure 42C D ( ). R. C. Harris Figure 42EF ( *). The whitish, sterile sorediate thallus containing stictic acid and coronatone characterize this species. Figure 42GH ( ). This material agrees with in the sorediate thallus and chemistry; however, the soralia are much smaller and discrete, resembling goniocystangia, and the thallus is greenish rather than whitish. Dibben ( ). DC. Figure 43AB ( *). Dibben Figure 43C ( *, *). Dibben Figure 43D ( *, ). (Fe) Nyl. Figure 43E ( *). Mll. Arg. Figure 43F ( ). R. C. Harris Figure 43G ( ). Mll. Arg. Figure 43H ( *, *, ). (Fink) Lendemer Figure 44A ( , *, ). Species of found in Florida are summarized in Table 6. (A. Massal.) Kalb & Matthes-Leicht. Figure 44BC ( , , ). Some of the specimens

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172 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Table 6. Keytable to identify species of and allied genera ( , ) cited in this work or otherwise reported from Florida. Septa = number of ascospore septa: muri = muriform; insp = hymenium inspersion; chem = secondary chemistry: pigm = quinone pigment (K+ green), lichx = lichexanthone; exci = excipulum carbonization; hypo = hypothecium (or basal part of excipulum) carbonization; lirellae = lirellae shape; disc = disc color and/or pruina; other = other characters: ecorticate = ecorticate thallus, lobulate = lobulate lirellae margin, orange = orange hymenium, striate = striate labia; size = ascospore size in m (typical upper range for length and size; the typical lower range is about 30% less). Species Septa Insp Chem Exci Hypo Lirellae Disc Other Size 5 pigm elongate-stellate dark red 30 8 3 norst elongate grey-pruinose 13 6 P. subfulgurata 3 norst elongate grey-pruinose lobulate 20 9 P. major 5 norst elongate grey-pruinose 22 8 3 norst pseudostromatic grey-pruinose 17 6 5 norst pseudostromatic grey-pruinose 22 6 3 stictic elongate brown (pruinose) striate 25 10 5 stictic pseudostromatic grey-pruinose 25 6 3 stictic stellate brown 25 9 P. aff. 5 elongate brown (pruinose) striate 30 10 5 insp lichx thin thin round-oval brown-black 25 8 3 insp norst oval-elongate brown-black 20 6 51 insp norst thin thin elongate-stellate brown-black ecorticate 40 10 P. lindigiana 3 insp elongate brown-black lobulate 18 8 3 insp thin thin elongate brown 25 7 5 insp thin thin elongate brown-black lobulate 30 8 51 insp thick thin round brown (pruinose) lobulate 40 10 muri norst elongate-stellate grey-pruinose 20 8 muri stellate grey-pruinose 35 12 muri insp norst thin elongate brown-black orange 35 10 muri insp elongate brown (pruinose) 100 30 P. aff. muri insp thin thin elongate brown (pruinose) 100 20 9 insp thick elongate bluish pruinose striate 70 13 muri insp norst thick elongate white-pruinose 35 10 muri insp thick elongate bluish pruinose 90 20 P. aff. muri insp thick elongate brown (pruinose) 90 20 3 insp thin thick stellate brown-black 18 7 5 insp thin thick elongate brown-black 30 8 muri insp thin thick elongate brown-black 28 11 3 insp stictic thin thick stromatic grey-pruinose 20 6 3 insp thin thin stellate (pruinose) 20 6

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 173 exhibited a negative K-reaction of thallus and lirellae, although TLC showed the presence of norstictic acid. Apparently, either the substance is concentrated in certain parts of the thallus or occurs in concentrations too low to be reliably detected by 10% KOH solution. The specimen supposedly devoid of lichen substances, in Harris (1995), was tested TLC negative and might represent a separate taxon. Specimens with stictic acid belong to P. *** Common & Lcking Figure 44D ( J, , ). See p. 148 for description of this new species. (Nyl.) Mll. Arg. Figure 44E ( *, ). Dal Forno & Eliasaro. Figure 44F ( , ). (Fe) Mll. Arg. Figure 44G ( ). This appears to be the correct name for the species commonly (Harris 1995). The latter is now considered a synonym of P. (Archer 2009). (Nyl.) Staiger Figure 44H ( ). (Kremp.) Mll. Arg. Figure 45A ( , ). This is the same taxon as in Harris (1995). There are two other species in Florida (not yet found at Fakahatchee) with small, muriform ascospores and norstictic acid, but with inspersed hymenium: R. C. Harris (with orange hymenium) and (Nyl.) Staiger, which in Harris (1995) keys out as There also seems to be an undescribed species from Florida with clear hymenium and smaller ascospores which requires further study. (Eschw.) Mll. Arg.Figure 45B ( ). (Kremp.) Lcking Figure 45C ( ). (Vain.) Zahlbr. Figure 45D ( ). (Ach.) Staiger Figure 45E ( ). aff. (Ach.) Staiger Figure 45F ( , ). We have not found a name for this taxon which resembles but has a completely carbonized excipulum. (Mll. Arg.) Mll. Arg. Figure 45G ( , , ). aff. (Mll. Arg.) Mll. Arg. Figure 45H ( , *, ). This material is the same as in Harris (1995). It agrees with in all details except the lack of lichen substances and larger ascospores. No name was found for this taxon, but revision of types of has not yet been concluded. (Nyl.) Zahlbr. ( ). spec. ( ). We have not found a name for this species. It is intermediate between in having much branched lirellae with noncarbonized base, 3-septate ascospores, and norstictic acid, and in having an inspersed hymenium. Swinscow & Krog Figure 46A ( , ). Zahlbr. Figure 46B ( ). (Vain.) Riddle Figure 46C ( , ). Brako Figure 46D ( *). Timdal Figure 46E ( ). This species was recently

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174 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) described from Peru and is also known from Cuba, Tobago, and Eduador (Timdal 2008), so its presence in subtropical Florida is not a surprise. It somewhat resembles but differs in thallus anatomy. (Tuck.) Swinscow & Krog. Figure 46F ( ). Moberg Figure 46G ( , *, *, ). Moberg Figure 46H ( ). (Fe) Fe Figure 47A ( ; , *, ). The species of are treated in Staiger (2002) and Tripp & Lendemer (2010). aff. (Fe) Fe Figure 47B ( , , ). This material differs from in the brown, thinly pruinose disc (bluish pruinose in ) and the usually pale to almost white thallus. (Fe) E. Tripp & Lendemer Figure 47C ( *). Not in checklist but reported by Tripp & Lendemer (2010). (Mll. Arg.) Staiger Figure 47D ( *, ). Not in checklist but reported by Tripp & Lendemer (2010). (Tuck.) Lendemer Figure 47E ( ). (Fe) Staiger Figure 47F ( ). (Kremp.) R. C. Harris Figure 47G ( ). (Nyl.) R. C. Harris ( ). (Nyl.) R. C. Harris Figure 47H ( *). (Fink J. Hedrick) R. C. Harris Figure 48A ( ). This species is similar to but differs in the larger ascospores with more numerous septa (9) which are distinctly tapering towards the proximal end; ascospores in are consistently 7-septate and symmetrically fusiform (Harris 1995). Ach. Figure 48BF ( , *, *, , *). T. H. Nash ( , , *). Mll. Arg. Figure 48G ( , ). Reinstated for the North American checklist. This taxon differs from (Fe) Mll. Arg. in the smaller ascospores (20 m versus 25 m) and the yellow inspersed hymenium (clear in ). The thallus is also more compact Mll. Arg. Figure 48H ( ). Reinstated for the North American checklist. This taxon differs from in the smaller ascospores and the largely endoperidermal thallus. (E. Michener) R. C. Harris ( ). aff. (E. Michener) R. C. Harris ( ). This material has longer ascospores with more numerous septa than typical P. (ascospores regularly 7-septate). spec. Figure 49AB ( , *). In spite of careful search we have not yet found a name for this diminutive but characteristic taxon. The thallus consists of very small (0.1 0.3 mm diam.), discrete, adnate squamules and bears abundant, orange-brown to red-brown apothecia with darker, persistent margin. Ascospores are non-septate or 1-septate and 12 1.5 m large, resembling those of Eventually the squamules break up to produce large, pale yellow-green soralia.

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 175 The taxon comes closest to (Nyl.) Mll. Arg. in the key given by Ekman (1996: 56), but that species has much longer ascospores and a -like prothallus and lacks soralia (Swinscow & Krog 1981). sorediate, microsquamulose Hepp (Printzen 1995), which differs in having -type apothecia with evanescent margin and much broader, ellipsoid ascospores. The since the type of that genus appears to belong in (S. Ekman, pers. comm. 2011). Henssen Figure 49CD ( ). (Ach.) Krog. ( ). R. C. Harris Figure 49E ( *, *). The genus is keyed out for Florida in Harris (1995). (Ach.) Vain. Figure 49F ( , , *). (Ach.) Ach. Figure 49G ( *, ). (Fe) R. C. Harris Figure 49H ( *, ). Fe Figure 50A ( ). (Nyl.) R. C. Harris Figure 50B ( *, , *). (Nyl.) R. C. Harris Synonym (Mll. Arg.) R. C. Harris Figure 50C ( ). (Mont.) Vain. Figure 50D ( , , *). (Mll. Arg.) R. C. Harris Figure 50E ( , *, *, ). (Borrer) Schaer. ( ). This species is characterized by a corticate, UV+ yellow thallus (lichexanthone), immersed perithecia covered mostly by a thalline layer except for the ostiolar area, non-inspersed hymenium, and ascospores 14 5 m with welldeveloped endospore and the terminal lumina separated from the exospore by an endospore layer. The species is regularly found in tropical montane situations and is also known from Europe; it is therefore surprising that it had not been reported before from North America. (Nyl.) Aptroot Figure 50F ( ). (Zahlbr.) R. C. Harris Figure 50G ( ). (Eschw.) Aptroot Figure 50H ( *). Ach. Figure 51A ( , , *, , *). (Ach.) Trevis. Figure 51B ( , *). R. C. Harris Figure 51C ( , ). (Vain.) R. C. Harris Figure 51D ( *, *, ). (Nyl.) R. C. Harris Figure 51E ( , , ). (Nyl.) R. C. Harris Figure 51F ( ). (Nyl.) Trevis. Figure 51G ( ). Fe Figure 51H ( , , ).

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176 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) (Nyl.) Mll. Arg. Figure 52A ( , , ). aff. (Nyl.) Mll. Arg. Figure 52B ( ). This material is the same as the one cited in Harris (1995: 106) for Collier County under the collection number It agrees with in the large perithecia covered by a thin thalline layer and the non-inspersed hymenium, but has smaller ascospores (14 4.5.5 m in the present material). We agree with Harris (1995) that this might represent a taxon distinct from s.str. (Eschw.) R. C. Harris ( *). (Nyl.) Mll. Arg. Figure 52C ( ). This taxon has almost invariably been included in a species with 3-septate ascospores (Harris 1995). Considering that ascospores with more than three transverse septa are extremely rare within and the material seen by us has either 3-septate or 5-septate mature ascospores, we propose to recognize the taxon with 5-septate ascospores as separate species. Tuck. Figure 52D ( ). (Mont. & Bosch) Nyl. Figure 52E ( , ). (Sw.) Nyl. Figure 52F ( ). (Tuck.) Vain. Figure 52G H ( *, , ). (Sw.) Ach. ( ). Tuck. ( *). Nyl. ( ). Nyl. ( ). De Not. ( ). Ach. ( ). Mll. Arg. ( , ). (L.) R. Howe ( , , *). R. Howe. ( ). (Tuck.) Kalb Figure 53A B ( *, ). (Ach.) Mll. Arg. Figure 53CH ( , , , , *, *, *). is possibly a collective species. The shape and arrangement of the lirellae within the stromata is very different across samples (Figure 53C H) and this appears to correlate with thallus morphology. (Ach.) Mll. Arg. Figure 54A ( , , ). (Durieu & Mont.) R.C. Harris Figure 54B ( *). Lcking & Lumbsch Figure 54C ( , , ). (Mont.) A. Frisch & Kalb ( ). (Kremp.) A. Frisch & Kalb Figure 54D ( ). In the checklist as (Tuck.) A. Frisch. Figure 54EH ( , , ). Delise ( ). ** A. L. Sm. Figure 55AB ( ). The genus was revised by Aptroot (2009), who stated that the

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 177 genus is almost absent from the Neotropics. Surprisingly, two species previously known only from the Paleotropics were found in our material, indicating that this genus is probably undercollected in the Neotropics or ** Makhija & Patw. Figure 55CD ( ). Fr. Figure 55E ( ). (Ach.) R. C. Harris Figure 55F ( , *). R. Sant. Figure 55G ( , ). Fr. Figure 55H ( , , ). (Vain.) Tehler Figure 56A ( *, *, ). Lcking Figure 56B ( , ). This species is described in a recent paper (Lumbsch et al. 2011) based on material from Costa Rica. It is anatomically similar to R. Sant. but differs in the distinct white pruina covering the apothecial margin. The ascospores are predominantly 3-septate but occasionally, ascospores with up American checklist, this name should replace which is with certainty only known from the Paleotropics. *** Common & Lcking Figure 56C ( ). See p. 149 for description of this new species. *** Lcking & Rivas Plata Figure 56D ( , *, ). See p. 149 for description of this new species. R. Sant. Figure 56E ( ). R. Sant. Figure 56F ( ). spec. Figure 56GH ( ). This material is characterized by a coarsely verrucose-bullate thallus; the hypothecium and excipulum are dark gray-brown reacting K+ sordid green, a reaction unusual for the genus (normally K+ purple). While campylidia are abundant in the material, only one apothecium was found and it lacked ascospores. Thus, while this taxon is certainly not identical with any known species of it cannot be formally described at this point, but further collections must be awaited to get ascospore data. (Hudson) Hafellner Figure 57A ( *). (Mont.) Trevis. Figure 57B ( , ). (Nyl.) Staiger Figure 57C ( *). Fe Figure 57D ( , , , *, *, ). The species is extremely variable in producing both perfectly round and distinctly lirellate ascomata on the same thallus (Harris 1995). Exactly the same variation was observed in material from the Philippines and the specimens cluster together in molecular phylogenetic analyses (Rivas Plata et al., unpubl. data). Tuck. Figure 57E ( , ). This species was synonymized with T. (Mll. Arg.) Mangold (Rivas Plata et al. 2010), but is actually set apart by the distinctly corticate thallus and extremely small apothecia. It frequently grows together with which has the same thallus type. Nyl. Figure 57F ( *, ). Mont. & Bosch. Figure

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178 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) 57G ( ). Tuck. ( ). Lcking & W. R. Buck. ( ). R. Sant. Figure 57H ( , ). (Eschw.) Zahlbr. Figure 58A ( , , *, *). Spreng. Figure 58B ( , *). (Fe) Aptroot Figure 58C Synonym (Zahlbr. Choisy) R. C. Harris( , , ). As in the North American checklist. (Nyl.) R. C. Harris Figure 58D ( , *, ). (Eschw.) Nyl. Figure 58E ( , *, *, , *). (Ach.) Mll. Arg. Figure 58F ( , , *, *). Nyl. Figure 58GH ( ). ACKNOWLEDGMENTS We thank the Fakahatchee Strand Preserve State Park staff and volunteers, in particular Mike Owen, Donna Glann-Smyth, Bob Nesmith, Alicia Campanella, Karen Relish, Laura Skinner, and Richard Fagan, for their support of this project and especially for their company and knowledgeable guidance through the park during our stay. The Florida Department of Environmental Protection, Division of Recreation and Parks, is kindly acknowledged for issuing collection permits in 1997 to Ralph Common (#04169714), in 2009 to William Safranek for the Tuckerman workshop (#4-09-02), and in 2010 to William Sanders (#410-34). Thanks also to Martha J. Robinson, Florida Department of Environmental Protection, for the permission to use the park map depicted in Figure 2. Linda Weinland organized the perfect laboratory facilities at Edison State College. Klaus Kalb of and and Andr Aptroot gave valuable advice regarding some of the pyrenocarpous taxa. The Tuckerman workshop was partially funded by the NSF project Neotropical Epiphytic Microlichens An Innovative Inventory of a Highly Diverse yet Little Known Group of Symbiotic Organisms (NSF-DEB 0715660 to The Field Museum; PI R. within the framework of phylogenetic studies of the lichen family Graphidaceae, supported by two NSF grants: Phylogeny and Taxonomy of Ostropalean Fungi (DEB 0516116; PI Lumbsch, Co-PI Lcking) and ATM Assembling a taxonomic monograph: The lichen family Graphidaceae (DEB 1025861; PI Lumbsch, Co-PI Lcking). Curators of the herbaria in B, BM, CANB, FH, G, H, M, NY, S, TUR, UPS, US, and W provided loans of, or access to, valuable type material and LITERATURE CITED Amtoft, A., F. Lutzoni, & J. Miadlikowska 2008. (Verrucariaceae) in the Ozark Highlands, North America. The Bryologist 111:1. Aptroot, A. 1991. A monograph of the Pyrenulaceae (excluding and ) and the Requienellaceae, with notes on

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LCKING ET AL. : Lichens of Fakahatchee Strand Preserve State Park 185 Peck, J. E., J. Grabner, D. Ladd, & D. R. Larsen. Ozarks lichens. The Bryologist 107:47. Perlmutter, G. B. 2006. in North America. The Bryologist 109:566. Petuch, E. J., & C. Roberts 2007. The Geology of the Everglades and Adjacent Areas. CRC Press, Taylor & Francis Group, Boca Raton, Florida. Pirozynski, K. A., & D. L. Hawksworth (eds.) 1988. Coevolution of Fungi with Plants and Animals. Academic Press, London. Printzen, C. 1995. Die Flechtengattung in Europa. Bibliotheca Lichenologica 60:1. Randazzo, A. F., & D. S. Jones (eds.) 1997. The Geology of Florida. University of Florida Press, Tallahassee. Reese, W. D., & S. Tucker. 1970. The 1967 foray of the American Bryological and Lichenological Society in Louisiana and Texas. The Bryologist 73:692. Rivas Plata, E., R. Lcking, A. Aptroot, H. J. M. Sipman, L. Umaa, J. L. Chaves, & D. Lizano biodiversity inventory in Costa Rica: the genus (Ostropales: Coenogoniaceae), with a world-wide key and checklist and a phenotype-based cladistic analysis. Fungal Diversity 23:255. Rivas Plata, E., R. Lcking, H. J. M. Sipman, A. Mangold, K. Kalb, & H. T. Lumbsch 2010. A world-wide key to the thelotremoid Graphidaceae, excuding the clade. The Lichenologist 42:187. Rivas Plata, E., J. E. Hernndez M., R. Lcking, B. Staiger, K. Kalb, & M. E. S. Cceres 2011. is two genera A remarkable case of parallel evolution in lichenized Ascomycota. Taxon 60 (in press). Robertson, W. B. Jr. 1955. An Analysis of Breedingbird Populations in Tropical Florida in Relation to the Vegetation. Ph.D. Dissertation, University of Illinois, Urbana, Illinois. Rolfs, P. H. 1901. Florida lichens. Transactions of the Academy of Sciences of St. Louis 11:25 39. Rosenzweig, M. L. 1995. Species Diversity in Space and Time. Cambridge University Press, Cambridge. Safranek, W. W., & Lcking, R. 2005. a new foliicolous lichen from the southeastern United States. The Bryologist 108:295. Schupp, E. W. 1992. The Janzen-Connell model for tropical tree diversity: population implications and the importance of spatial scale. American Naturalist 140:526. Seavey, F. 2009. sp. nov. (Arthoniaceae), a new lichen species from Everglades National Park, Florida. Opuscula Philolichenum 7:49. Seavey, F., & J. Seavey. 2009. Subtropical Florida Lichens/default.htm (accessed May 2009). Srusiaux, E. 1979. Foliicolous lichens from southeastern United States. The Bryologist 82:88. Shmida A., & E. O. Wilson 1985. Biological determinants of species diversity. Journal of Biogeography 12:1. Sipman, H. J. M., & A. Aptroot 2001. Where are the missing lichens? Mycological Research 105:1433. Skorepa, A. C. 1968. Lichens from the Okefenokee Swamp, Georgia. Castanea 33:248. Smith, C. W. 1986. Three foliicolous lichens new to the United States. The Bryologist 89:232. Sparrius, L. B. & W. Saipunkaew. 2005. a new species from northern Thailand. Lichenologist 37:507 509. Spribille, T., S. Prez-Ortega, S., T. Tnsberg, & D. Schirokauer 2010. Lichens and lichenicolous fungi of the Klondike Gold Rush National Historic Park, Alaska, in a global biodiversity context. The Bryologist 113:439. Staiger, B., & K. Kalb. 1999. and other graphidioid lichens with warty periphysoids or paraphysis-tips. Mycotaxon 73:69. Staiger, B. 2002. Die Flechtenfamilie Graphidaceae. Studien in Richtung einer natrlicheren

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186 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 49(4) Gliederung. Bibliotheca Lichenologica 85:1 526. Staiger, B., K. Kalb, & M. Grube 2006. Phylogeny and phenotypic variation in the lichen family Graphidaceae (Ostropomycetidae, Ascomycota). Mycological Research 110:765 772. Stocker, G. C., G. L. Unwin, & P. W. West 1985. Measures of richness, evenness and diversity in tropical rainforest. Australian Journal of Botany 33:131. Swinscow, T. D. V., & H. Krog 1981. The genus with a report on the East African species. The Lichenologist 13:203. Thomson, J. W. 2003. Lichens of Wisconsin. Wisconsin State Herbarium, Department of Botany, University of Wisconsin, Madison. Thor, G. 1988. Three calicialian lichens new to the continental United States. The Bryologist 91:367. Timdal, E. 2008. Studies on (Ramalinaceae) in Peru. The Lichenologist 40:337. Tomlinson, P. B. 1980. The Biology of Trees Native to Tropical Florida, 2 nd Edition. Published by the author, printed by the Harvard University Tripp, E. A. & J. C. Lendemer. 2010. The genus in North America. Castanea 75:388. Tripp, E. A., J. C. Lendemer, & R. C. Harris. 2010. Resolving the genus Mll. Arg. in North America: new species, new combinations, and treatments for , and Lichenologist 42:55. Tucker, S. C. 1979. New or noteworthy records of lichens from Louisiana. The Bryologist 82:125. Tucker, S. C. 1981. Checklist of Louisiana lichens. Proceedings of the Louisiana Academy of Sciences 44:58. Tucker, S. C. 2010. Lichens of Burden Research Plantation, Baton Rouge, Louisiana. Evansia 27:121. Tucker, S. C., & R. C. Harris. 1980. New or noteworthy pyrenocarpous lichens from Louisiana and Florida. The Bryologist 83:1 20. Tuckerman, E. 1872. Genera Lichenum: an arrangement of the North American lichens. Amherst. Tuckerman, E. 1888. A synopsis of the North American lichens. Part. II. Comprising the Lecideacei, and (in part) the Graphidacei. New Bedford, Massachussetts. Waterman, P. G., & D. McKey. 1989. Herbivory and secondary compounds in rainforest plants. Pp. 513 H. Lieth & M. J. A. Werger (eds.). Tropical Rain Forest Ecosystems. Biogeographical and Ecological Studies [Ecosystems of the World 14B]. Elsevier, Amsterdam. Wetmore, C. M. 1994. The lichen genus in North and Central America with brown or black apothecia. Mycologia 86:813. Wills, C., R. Condit, R. B. Foster, & S. P. Hubbell. 1997. Strong densityand diversity-related effects help to maintain tree species diversity in a neotropical forest. Proceedings of the National Academy of Sciences of the USA 94:1252. Wirth, M., & M. E. Hale Jr. 1978. MordenSmithsonian Expedition to Dominica: the lichens (Graphidaceae). Smithsonian Contributions to Botany 40:1.

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The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY publishes original research conducted by our faculty, staff, students, and research associates. We also accept submissions of appropriate, fully funded manuscripts from external researchers. Priority is given to monograph-length papers describing be exceptions as determined by the Managing Editor. Starting in 2010, the Bulletin is published simultaneously in two formats. Approximately 400 printed copies are distributed to libraries and museums world-wide by the University of Florida Library system. Authors have the option of purchasing additional printed copies at cost for distribution to colleagues and associates. An identical, electronic version is posted in PDF format on the Florida Museum of Natural History web site simultaneously with the availability of the printed version and is available free of charge for reading or downloading. Supplemental materials are available only through the web site. INSTRUCTIONS FOR AUTHORS papers must adhere to the rules published in the appropriate international code of systematic nomenclature. RECENT PUBLICATIONS OF THE FLMNH BULLETIN Thomson, A. W., & L. M. Page. 2010. Taxonomic revision of the Franz, R., & S. E. Franz. 2009. A new fossil land tortoise in the genus Zaspel, J. M., S. J. Weller, & R. T. Carde. 2008. A review of $7.00 16. $7.00 A complete list of available issues and current prices of the Bulletin of the Florida Museum of Natural History

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Figure 1. Group photograph of the 2009 Tuckerman workshop. Standing from left to right: Lee Crane, Sean Beeching, Frederick Seavey, Ma lcolm Hodges, Richard Fagan, William Buck, Elisabeth Lay, Mike Owen, Harold Schaefer, Matthew Nelsen, William Sanders, Troy McMullin, and Brendan Hodkinson. Kneeling from left to right: Jean Seavey, Bob Nesmith, Alicia Campanella, James Lendemer, William Sa franek, Ottmar Breuss, Joel Mercado-Daz, Eimy Rivas Plata, Natasha Lcking Rivas Plata, and Robert Lcking. Figure 2. A, County map of Florida showing Collier County and the study site. B, Map of Fakahatchee Strand Preserve Stat e Park, straddling the Tamiami Trail (Route 41) within Collier County; the four main collection sites described in the text are indicated by numerals 1 to 4. Map reprinted with permission of the Florida Depa rtment of Environment Protection, Tallahassee, Florida. From on-line supplemental materials to: Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18 th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):1276.

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Figure 3. Habitat photographs of the four collecti on sites at Fakahatchee Strand Preserve State Park. First row: site 1, showi ng royal palm canopy and understory with bald cypress. Second row: site 2, showing side road along old tram with dense vegetation a nd aspect of understory with bromeliads. Third row: site 3, showing open bald cypress prairie hammock and dense regrowth. Fourth row: site 4, show ing closed bald cypress forest and aspect of forest understory. Photographs by R. Lcking (A, B, G), Matthew Nels en (D, E, H), Frederick Seavey (F), and Jean Seavey (C).

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From on-line supplemental materials to: Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18 th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):1276.

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SUPPLEMENTAL MATERIALS FIGURES 4 58 Lcking, R., F. Seavey, R. S. Common, S. Q. Beeching, O. Breuss, W. R. Buck, L. Crane, M. Hodges, B. P. Hodkinson, E. Lay, J. C. Lendemer, R. T. McMullin, J. A. Mercado Daz, M. P. Nelsen, E. Rivas Plata, W. Safrane k, W. B. Sanders, H. P. Schaefer Jr. & J. Seavey. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127186 Copyright for individual images belongs to the listed photographer.

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Figure 4. A B Acanthothecis peplophora (A, Lcking 26501 ; B, Seavey & Seavey 11007 ). C Acanthothecis poitaeoides ( Seavey & Seavey 10843 ). D Actinoplaca spec. ( Sanders 10504.6). E Amandinea endachroa ( Bre uss 28936). F Amandinea punctata ( Seavey & Seavey 10712 ). G Anisomeridium subnexum ( Lcking & Rivas Plata 26616). H Anisomeridium subprostans ( Seavey & Seavey 10700 ). Photographs R. Lcking (A B, D E, G), R. Seavey (C, F, H). From on line s upplement al m aterials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 5. A Anisomeridium spec. ( Lck ing & Rivas Plata 26598 ). B D Anthracothecium prasinum ( Lcking & Rivas Plata 26595, 26594, 26292). E F, Arthonia antillarum ( Common 7327E ). G H, Arthonia cinnabarina ( Common 7421C ) Photographs R. Common (E H), R. Lcking (A D). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 6. A Arthonia complanata ( Se avey & Seavey 10542 ). B Arthonia ilicina ( Mercado Daz 410 ). C D Arthonia interveniens ( Common 7382A ). E F, Arthonia macrotheca ( Seavey & Seavey 10167 ). G H, Arthonia mirabilis ( Common 7265 ). Photographs R. Common (C D, G H), R. Lcking (B), R. Seavey (A E F). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Fig ure 7. A Arthonia aff. pinastri ( Lay 09 0042 ). B Arthonia platygraphidea ( Lcking & Rivas Plata 26583 ). C D Arthonia aff. pruinata ( Common 7372A ). E F, Arthonia pruinosella ( Common 7327A ). G H, Arthonia pyrrhuliza ( Lay 09 0017A ). Photographs R. Common ( C F), R. Lcking (A B, G H). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 4 9(4):127 186.

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Figure 8. A B Arthonia redingeri ( Common 7259U ). C Arthonia rubella ( Lcking & Rivas Plata 26586). D Arthonia rubrocincta ( Lcking & Rivas Plata 26582 ). E F, Arthonia simplicascens ( Common 7291D ). G H, Arthonia speciosa ( Common 7291I ). Photographs R. Common (A B, E H), R. Lcking (C D). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Mus eum of Natural History 49(4):127 186.

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Figure 9. A Arthopyrenia cinchonae B Arthopyrenia lyrata. C Arthopyrenia planorbis ( Seavey & Seavey 10165 ). D Arthopyrenia aff. planorbis ( Lcking & Rivas Plata 26787b ). E Arthothelium spectabile ( Breuss 2877 0 ). F, Aspidothelium cinerascens G, Aspidothelium geminiparum ( Lcking & Rivas Plata 26600c ). H Aspidothelium scutellicarpum ( Breuss 28787 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Faka hatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 10. A Astrothelium confusum ( Lcking & Rivas Plata 26691). B Astrothelium diplocarpoide s ( Lcking & Rivas Plata 26627). C Astrothelium diplocarpum ( Nelsen ). D Astrothelium galbineum ( Lcking & Rivas Plata 26596). E F, Astrothelium variolosum ( Breuss 28962 ). G, Aulaxina microphana ( Sanders 10521.1). H Aulaxina quadrangula hyphophores ( Sanders 10521.7). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Nat ural History 49(4):127 186.

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Figure 11. A Bacidia aggregatula ( Seavey & Seavey 10095 ). B Bacidia campalea ( Breuss 28735 ). C Bacidia heterochoa ( Lcking & Rivas Plata 26621 ). D Bacidia aff. heterochroa ( Lcking & Rivas Plata 26610 ). E Bacidia hosthe leoides ( Lcking & Rivas Plata 26605 ). F Bacidia aff. hostheleoides ( Lcking & Rivas Plata 26600b ). G Bacidia medialis ( Lcking & Rivas Plata 26601 ). H, Bacidia aff. medialis ( Lcking & Rivas Plata 26606). Photographs R. Lcking (B H), R. Seavey (A). Fro m on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 12. A Bacidia mutabilis ( Lcking & Rivas Plata 26603 ). B Bacidia russeola ( Breuss 28828 ). C Bacidia schweinitzii ( Seavey & Seavey 13331 ). D Bacidia aff. schweinitzii ( Lcking & Rivas Plata 26602a ). E Bacidia spec. ( Lcking & Rivas Plata 26611 ). F, Bacidina v aria ( Lay 09 0004). G, Bactrospora denticulata ( Lcking & Rivas Plata 26737b ). H Bactrospora myriadea ( Lay 09 0212). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve S tate Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 13. A Baculifera curtisii ( Seavey & Seavey 10432 ). B Baculifera imshaugiana ( Breuss 28890 ). C Bathelium carolin ianum ( Nelsen 4149 ). D Bathelium madreporiforme ( Lay 09 0047 ). E Brigantiaea leucoxantha ( Breuss 28981 ). F, Byssoloma chlorinum ( Nelsen ). G, Byssoloma leucoblepharum ( Lcking & Rivas Plata 26692 ). H, Byssoloma meadii ( Lendemer 15572). Photographs R. Lcking (B H), R. Seavey (A). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4 ):127 186.

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Figure 14. A B Calicium hyperelloides ( Seavey & Seavey). C Calopadia editae ( Common 7322B ). D G, Calopadia floridana (D E, holotype; F G, Common 5320K ). H, Calopadia fusca ( Seavey & Seavey 10365 ). Photographs R. Common (C, F G), R. Lcking (D E), R. Seavey (A B, H). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49 (4):127 186.

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Figure 15. A F, Calopadia imshaugii (A B, E, holotype; C D, Homestead, Common 5889I ; F, Homestead, Common 5892E ). G H, Calopadia lecanorella ( Nelsen s.n. Lcking & Rivas Plata 26696 ). Photographs R. Common (A F), R. Lcking (G H). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 16. A Calo padia perpallida ( Nelsen ). B Calopadia puiggarii C Calopadia subcoerulescens D Caloplaca camptidia ( Lcking & Rivas Plata 26568b ). E Caloplaca epiphora ( Seavey & Seavey 10543 ). F, Caloplaca holocarpa ( Lcking & Rivas Plata 26698). G, Catillochroma en dochroma ( Beeching ). H, Celothelium aciculiferum ( Common 7396D ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerm an Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 17. A Chaenotheca brunneola ( Lay 09 0049 ). B Chapsa chionostoma ( Common 7321A ). C Chapsa platycarpa ( Lcking & Rivas Plata 26573 ). D Chapsa platycarpoides ( Lcking & Rivas Plata 26571 ). E Chrysothrix xanthina ( Nelsen ). F, Clathroporina isidiifera ( Lcking & Rivas Plata 26768 ). G, Clathroporina subpungens ( Seavey & Seavey 10241 ). H, Clathroporina tetracerae ( Lcking & Rivas Plata 26767 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 18. A Coccocarpia domingensis B Coccocarpia erythroxyli ( Seavey & Seavey 10382 ). C D Coccocarpia palmicola ( Lcking & Rivas Plata 26701 ). E Coenogonium congense ( Lay 09 0134). F, Coenogonium geralense ( Lcking & Rivas Plata 26702). G Coenogonium interple xum ( Lay 09 0132 ). H, Coenogonium linkii ( Lay 090002 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 19. A Coenogonium luteocitrinum ( Lay 09 0131 ). B Coenogonium luteum C Coenogonium subdentatum ( Lay 09 0130). D Coenogonium subfallaciosum ( Lcking & Rivas Plata 26703 ). E Coniarthonia wilmsiana ( Common 7291G). F Cratiria lauricassiae ( Common 7287E ). G, Crocynia gossypina ( Seavey & Seavey 10473 ). H, Crocynia pyxinoides ( Lcking & Rivas Plata 26705 ). Photographs R. Common (E), R. Lcking (all others). From on line supplemen tal materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 20. A Cryptolechia nana ( Breuss 28711 ). B Cryptothecia effusa C D Cryptothecia evergladensis ( Lcking & Rivas Plata 26670). E F, Cryptothecia miniata ( Lcking & Rivas Plata 26678 ). G Cryptothecia punctosorediata ( Lcking & Rivas Plata 26680 ). H, Cryptothecia striata ( Lcking & Rivas Plata 26687 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 22. A B Dirinaria papilluli fera (Colombia, Lcking 32504). C Dirinaria picta ( Seavey & Seavey 10633 ). D Dirinaria purpurascens ( Seavey & Seavey 10636 ). E F, Dyplolabia afzelii ( Lcking & Rivas Plata 26509, 26505). G, Echinoplaca areolata ( Common 7285B ). H, Echinoplaca aff. leucotr ichoides ( Crane ). Photographs R. Lcking (A B, D H), R. Seavey (C). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 23. A Echinoplaca lucernifera ( Sanders 10504.1b ). B Echinoplaca similis ( Breuss 28922 ). C Enterographa anguinella ( Seavey & Seavey 10429 ). D Eugeniella leucocheila E Fissurina aggregatul a (holotype). F G, Fissurina analphabetica (holotype). H, Fissurina cingalina ( Common 7323F ). Photographs R. Common (G), R. Lcking (all others). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve S tate Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 24. A Fissurina columbina ( Lendemer ). B Fissurina confusa ( Common 7356A ). C D Fissurina crassilabra ( Common 735 6C ). E Fissurina egena ( Common 7276B 7368Q ). F Fissurina aff. elaiocarpa ( Common 7356T ). G H, Fissurina humilis (Homestead, Common 5887F ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 25. A Fissurina illiterata ( Lcking & Rivas Plata 26618 ). B Fissurina inspersa (holotype ). C Fissurina mexicana ( Lcking & Rivas Plata 26513). D Fissurina pseudostromatica (holotype). E Fissurina radiata ( Common 7356A ). F, Fissurina rufula ( Breuss 28769 ). G Fissurina subcomparimuralis (holotype). H, Fissurina subnitidula ( Common 7418H ). A ll photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49( 4):127 186.

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Figure 26. A Fissurina tachygrapha ( Common 7418A ). B Fissurina tuckermaniana (holotype). C Fissurina varieseptata (holotype). D Flakea papillata ( Lay 09 0062 ). E Glyphis atrofusca ( Lay 09 0015). F G Glyphis cicatricosa ( Common 7259C ). H, Glyphis scyphulifera ( Common 7292D ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 27. A Graphis cf. acharii ( L cking & Rivas Plata 26521). B Graphis anfractuosa ( Lay 09 0070 ). C Graphis aperiens ( Common 7425D ). D E Graphis appendiculata (D, holotype; E, Lcking & Rivas Plata 26657 ). F, Graphis argentata ( Breuss 28825 ). G, Graphis assimilis (Everglades City, Common 7346I ). H, Graphis caesiella (Everglades City, Common 7341A ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 28. A Graphis caesiocarpa ( Breuss 28718 ). B Graphis caribica ( Common 7356G ). C Graphis conferta ( Lay 09 0016 ). D Graphis cupei ( Lcking & Rivas Plata 26536a). E Graphis desquamescens ( Common 7368O ). F G Graphis disserpens ( Lcking & Rivas Plata 26656 ). H, Graphis handelii ( Common 7292G ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 29. A Gr aphis longula ( Lcking & Rivas Plata 26527 ). B Graphis lucifica ( Lay 09 0065). C Graphis oshioi ( Lcking & Rivas Plata 26612 ). D Graphis oxyclada ( Hodges s.n. ). E G Graphis pseudocinerea Lcking (E, Breuss 28987 ; F G, Lcking & Rivas Plata 26537 ). H, G raphis rimulosa (Costa Rica: Standley 71849 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bullet in of the Florida Museum of Natural History 49(4):127 186.

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Figure 30. A Graphis sauroidea ( Breuss 28890 ). B Graphis subflexibilis ( Common 7380G ). C F, Graphis stellata ( Breuss 28889 ). G, Graphis xanthospora ( Lcking & Rivas Plata 26535). H, Graphis x ylophaga ( Seavey & Seavey 10359). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Fl orida Museum of Natural History 49(4):127 186.

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Figure 31. A B Gyalectidium appendiculatum ( Sanders 10504.6). C Gyalectidium catenulatum ( Sanders 10504.8). D E Gyalectidium floridense ( Sanders 10521.4 ). F, Gyalectidium imperfectum ( Sanders 10504.7 ). G Gyalectidium ulloae ( Sanders 10504.10). H, Gyalectidium aff. yahriae ( Sanders 10521.2). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceed ings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 32. A Haematomma flexuosum ( Nelsen ). B Haematomma guyanense C Haematomma persoonii ( Seavey & Seavey 10070 ). D E Hafellia bahiana (Evergl ades: Seavey & Seavey ). F Hafellia curatellae ( Seavey & Seavey 10615). G, Hafellia pleiotera ( Breuss 28936). H, Halegrapha floridana ( Common 7410C ). Photographs R. Lcking (A B, G H), R. Seavey (C F). From on line supplemental materials to : Lcking, R., e t al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 33. A B Heiomasia seaveyorum C D Herpothallon antillaru m ( Nelsen 4037). E Herpothallon echinatum ( Lcking & Rivas Plata 26681 ). F G Herpothallon rubrocinctum (F, Lcking & Rivas Plata 26707; G, Lcking & Rivas Plata 26708 ). H, Herpothallon rubroechinatum ( Lcking & Rivas Plata 26682). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 34. A Heterodermia albicans ( Breuss 28763 ). B Heterodermia pseudospeciosa ( Breuss 29020 ). C D Laurera megasperma (C, Lcking & Rivas Plata 26709; D, Lcking & Rivas Plata 26710a). E Lecanora achroa ( Seavey & Seavey 10144 ). F, Lecanora achroides ( Seave y & Seavey 10624 ). G Lecanora allophana ( L cking & Rivas Plata 26639 ). H, Lecanora argentata ( Lcking & Rivas Plata 26634b). Photographs R. Lcking (A D, G H), R. Seavey (E F). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 35. A Lecanora caesiorubella ssp. glaucomodes ( Seavey & Seavey 10077 ). B C Lecan ora floridula ( Lay 09 0083 ). D Lecanora hybocarpa ( Lay 09 0138). E Lecanora leprosa ( Breuss 28936 ). F, Lecanora pseudargentata ( Lcking & Rivas Plata 26642 ). G, Lecanora strobilina ( Lcking & Rivas Plata 26637 ). H Leiorreuma exaltatum ( Common 7424A ). Ph otographs R. Lcking (B H), R. Seavey (A). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Nat ural History 49(4):127 186.

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Figure 36. A Leiorreuma sericeum ( Common 7355E ). B Letrouitia domingensis ( Lcking & Rivas Plata 26718). C D Letrouitia vulpina (C, Lcking & Rivas Plata 26723; D, Nelsen ). E Leucodecton compunctellum ( Lcking & Rivas Pl ata 26566b ). F G Leucodecton glaucescens ( Nelsen) H, Leucodecton occultum ( Common 7321B ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Procee dings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 38 A B Megalospora porphyritis ( Hodges s.n.) C Micarea prasina ( Lay 09 0088 ). D G Micarea spec. ( Lay 090051). H, Mycocalicium subtile ( Lay 09 0097 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The liche ns of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 39. A Mycomicrothelia modesta ( Lcking & Rivas Plata 26588a). B Mycomicrothel ia subfallens ( Lcking & Rivas Plata 26772 ). C D Mycomicrothelia willeyana ( Lcking & Rivas Plata 26733). E F, Mycoporum lacteum ( Lcking & Rivas Plata 26732 ). G H, Mycoporum sparsellum ( Lcking & Rivas Plata 26613 ). All photographs R. Lcking. From on li ne supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 40. AB Myriotrema erodens (A, Harris 29645; B, Lcking & Rivas Plata 26541 ). C Myriotrema peninsulae ( Lcking & Rivas Plata 26542 ). D Myriotrema pycnoporellum ( Lcking & Rivas Plata 26544 ). E Nadvornikia hawaiiensis F Nadvornikia sorediata G Ocellularia auberianoides ( Lcking & Rivas Plata 26548 ). H, Ocellularia obturascens ( Mercado-Daz 381 ). Photographs R. Lcking (BH), A. Mangold (A). From on-line supplemental materials to: Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18 th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):1276.

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Figure 41. A Ochrolechia africana ( Seavey & Seavey 10752 ). B C Opegrapha astraea ( Lcking & Rivas Plata 26734). D Opegrapha atra ( Lckin g & Rivas Plata 26737a ). E Opegrapha candida ( Seavey & Seavey 10628 ). F, Opegrapha cypressi ( Seavey & Seavey 10791 ). G, Opegrapha longissima ( Lcking & Rivas Plata 26815 ). H, Opegrapha spec. ( Mercado Daz 450). Photographs R. Lcking (A E, G H), R. Seavey (F). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figur e 42 A B Parmeliella stylophora ( Lcking & Rivas Plata 26738). C D Pertusaria epixantha ( Lcking & Rivas Plata 26666c ). E F Pertusaria expolita ( Lcking & Rivas Plata 26650 ). G H, Pertusaria aff. expolita ( Lcking & Rivas Plata 26649 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 43. A B Pertusaria leioplaca ( Lcking & Rivas Plata 26623c ). C Pertusaria paratuberculifera ( Lcking & Rivas Plata 26645 ). D Pertusaria sinusmexicani ( Lcking & Rivas Plata 26654 ). E Pertusaria tetrathalamia F, Pertusaria texana ( Lcking & Riva s Plata 26643 ). G Pertusaria virensica ( Lay 09 0141). H Pertusaria xanthodes ( Lcking & Rivas Plata 26647 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Par k, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 44. A Phaeographis asteroides ( Lay 09 0145). B C Phaeographis brasiliensis (B, Brazil: Cceres 874 ; C, Common 7355M ). D Phaeographis delicatula (holotype). E Phaeographis erumpens ( Common 7277A ). F, Phaeographis flavescens ( Common 7355K ). G Phaeographis inconspicua ( Lay 090149). H Phaeographis intricans ( Common 7355A ). All photographs R. Lcking. From on line supplemen tal materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 45. A Phaeographis leiogr ammodes ( Common 7277C ). B Phaeographis lobata (Homestead, Common 5891A ). C Phaeographis major ( Common 7355B ). D Phaeographis nylanderi ( Common 7346A ). E Phaeographis scalpturata ( Seavey & Seavey 10297 ). F, Phaeographis aff. scalpturata ( Lcking & Rivas Plata 26740 ). G Phaeographis schizoloma ( Common 7355C ). H, Phaeographis aff. schizoloma ( Lcking & Rivas Plata 26746). Photographs Ralph Common (B), R. Lcking (all others). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 46. A Phyllopsora confusa ( Nelsen ). B Phyllopsora furfuracea. C Phyllopsora isidi otyla ( Lcking & Rivas Plata 26748 ). D Phyllopsora kalbii ( Lcking & Rivas Plata 26750). E Phyllopsora lacerata ( Sanders s.n. ). F, Phyllopsora santensis (Lay 09 0211). G Physcia atrostriata (F, Lcking & Rivas Plata 26753; G, Lcking & Rivas Plata 26751 ). H, Physcia undulata. All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Muse um of Natural History 49(4):127 186.

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Figure 47. A Platygramme caesiopruinosa ( Lcking & Rivas Plata 26826b ). B Platygramme aff. caesiopruinosa ( Common 7259E ). C Platygramme pachnodes ( Seavey & Seavey 10300 ). D Platygramme praestans ( Lcking & Rivas Plata 26628). E Platythecium floridanum ( Common 7368P ). F, Platythecium grammitis (Costa Rica, Lcking 17100a). G Polymeridium albocinereum ( Lay 09 0143). H, Polymeridium proponens ( Seavey & Seavey 10302 ). All photographs R. Lcking. From on line supple mental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 48. A Porina heterospora B F Porina nucula (B, Lcking & Rivas Plata 26760; C, Lcking & Rivas Plata 26761 ; D Lcking & Rivas Plata 26762; E, Lcking & Rivas Plata 26763; F, Lcking & Rivas Plata 26766 ). G Pseudopyrenula subgregaria ( Lcking & Rivas Plata 26770). H, Pseud opyrenula subnudata (Costa Rica, Lcking s.n. ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bull etin of the Florida Museum of Natural History 49(4):127 186.

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Figure 49. A B Psorella spec. ( Lcking & Rivas Plata 26829 ). C D Psoroglaena costaricensis ( Common 7363D). E Pyrenula acutalis ( Breuss 28797 ). F, Pyrenula anomala ( Lcking & Rivas Plata s.n. ). G, Pyrenula aspistea ( Breuss 28679 ). H, Pyrenula astroidea ( Lcking & Rivas Plata 26774 ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: P roceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 50. A Pyrenula brunnea ( Lcking & Rivas Plata 26777 ). B Pyrenula concatervans ( Lcking & Rivas Plata 26778). C Pyrenula confinis ( Co mmon 7281G ). D Pyrenula cruenta ( Lcking & Rivas Plata 26781). E Pyrenula cubana ( Lcking & Rivas Plata 26783 ). F Pyrenula duplicans ( Lcking & Rivas Plata 26785a). G, Pyrenula falsaria ( Lcking & Rivas Plata 26786). H, Pyrenula globifera ( Breuss 28709 ) All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 51. A Pyrenula leucostoma ( Lcking & Rivas Plata 26789). B Pyrenula mamillana ( Lcking & Rivas Plata 26782). C Pyrenula microtheca ( Common 7326B ). D Pyrenula mucosa ( Lcking & Rivas Plata 26812). E Pyrenula ochraceoflava ( Sea vey & Seavey 10309 ). F Pyrenula ochraceoflavens ( Seavey & Seavey 10311 ). G Pyrenula punctella ( Lcking & Rivas Plata 26801). H, Pyrenula quassiicola ( Mercado Daz 432). Photographs R. Common (C), R. Lcking (A B, D, G H), R. Seavey (E F). From on line su pplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 52. A Pyrenula san tensis ( Lcking & Rivas Plata 26785 ). B Pyrenula aff. santensis ( Mercado Daz 442 ). C Pyrenula sexlocularis ( Lcking & Rivas Plata 26780). D Pyrenula thelomorpha ( Lcking & Rivas Plata 26658b). E Pyrgillus javanicus ( Lay 09 0110). F Pyxine cocoes ( Lc king & Rivas Plata 26806 ). G H Pyxine eschweileri (A, Lcking & Rivas Plata 26563b; B, Lay 09 0060). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Flori da: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 53. A B Reimnitzia santensis (A, Nelsen ; B, Lcking & Rivas Plata 26558). C H Sarcographa labyrinthica (C D, Lcking & Rivas Plata 26559 ; E F Common 7367A ; G H Common 7381A ). All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bul letin of the Florida Museum of Natural History 49(4):127 186.

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Figure 54. A Sarcographa tricosa ( Common 7346H ). B Segestria leptalea C Sporopodium marginatum ( Lcking & Rivas Plata 26808a ). D Stegobolus emersus E H, Stegobolus granulosus ( Nelsen) All photographs R. Lcking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 4 9(4):127 186.

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Figure 55. A B Stirtonia dubia ( Common 7421D ). C D Stirtonia macrocarpa ( Common 7327C ). E Strigula orbicularis ( Seavey & Seavey 10510 ). F, Strigula phaea ( Lcking & Rivas Plata 26589). G, Strigula schizospora ( Lay 09 0208). H, Strigula smaragdula ( Lay 09 0208 ). Photographs Ralph Common (A D), R. Lcking (F H), R. Seavey (E). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerma n Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 56. A Syncesia byssina ( Lcking & Rivas Plata 26831). B Tapellaria albomarginata (Costa Rica, Crane 195). C Tapellaria floridensis (holotype). D Tapellaria granulos a (holotype). E Tapellaria malmei ( Lcking & Rivas Plata 26811 ). F, Tapellaria nana ( Seavey & Seavey 10618 ). G H, Tapellaria spec. ( Hodges ). Photographs R. Lcking (A E, G H), R. Seavey (F). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 57. A Tephromela atra ( Seavey & Seavey 10574 ). B Thalloloma anguinum ( Common 7367B ). C Thalloloma hypoleptum ( Lcking & Rivas Plata 26564 ). D Thecaria quassiicola ( Lcking & Rivas Plata 26626 ). E Thelotrema lathraeum ( 7371A ). F, Thelotrema pachysporum ( Lcking & Rivas Plata 26569). G, Thelotrema porinoides ( Lcking & Rivas Plata 26570). H, Tricharia vainioi ( Lcking & Rivas Plata 26808c ). Photographs R. Lcking (B H), R. Seavey (A). From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceed ings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.

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Figure 58. A Trypethelium aeneum ( Lcking & Rivas Plata 26814 ). B Trypethelium eluteriae ( Seavey & Seavey 10532). C Trypethelium marcidum ( Merca doDaz 479 ). D Trypethelium nitidiusculum ( Lcking & Rivas Plata 26816). E Trypethelium ochroleucum ( Mercado Daz 417 ). F Trypethelium tropicum ( Lcking & Rivas Plata 26827). G H, Tylophoron moderatum (Costa Rica, Lcking 15081a). All photographs R. L cking. From on line supplemental materials to : Lcking, R., et al. 2011. The lichens of Fakahatchee Strand Preserve State Park, Florida: Proceedings from the 18th Tuckerman Workshop. Bulletin of the Florida Museum of Natural History 49(4):127 186.