Studies in Lichens and Lichenicolous Fungi – No. 22: The identities of Lecidea deminutula, L. olivacea var. inspersa, L. virginiensis and Thelenella humilis

Abstract

Notes on four taxa are presented as part of an effort to resolve the taxonomic status of neglected crustose lichen names based on material from North America. Lecidea deminutula, described from non-calcareous rocks in the Great Smoky Mountains of Tennessee, is placed in synonymy with Lecidella enteroleucella. Lecidea olivacea var. inspersa, described from hardwood bark in the Great Smoky Mountains of Tennessee, is placed in synonymy with Lecidella elaeochroma. Lecidea virginiensis, described from seeping, non-calcareous rocks in West Virginia, is lectotypified and placed in synonymy with Bryobilimbia ahlesii. Thelenella humilis described from non-calcareous rocks in New York is placed in synonymy with Protothelenella corrosa.

Full text

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Opuscula Philolichenum, 18: 90-101. 2019.
*pdf effectively published online 7August2019 via (http://sweetgum.nybg.org/philolichenum/)
Studies in Lichens and Lichenicolous Fungi – No. 22: The
identities of Lecidea deminutula, L. olivacea var. inspersa,
L. virginiensis and Thelenella humilis
JAMES C. LENDEMER
1
, RICHARD C. HARRIS
2
AND R. TROY MCMULLIN
3
ABSTRACT. – Notes on four taxa are presented as part of an effort to resolve the taxonomic
status of neglected crustose lichen names based on material from North America. Lecidea deminutula,
described from non-calcareous rocks in the Great Smoky Mountains of Tennessee, is placed in synonymy
with Lecidella enteroleucella. Lecidea olivacea var. inspersa, described from hardwood bark in the Great
Smoky Mountains of Tennessee, is placed in synonymy with Lecidella elaeochroma. Lecidea virginiensis,
described from seeping, non-calcareous rocks in West Virginia, is lectotypified and placed in synonymy
with Bryobilimbia ahlesii. Thelenella humilis described from non-calcareous rocks in New York is placed
in synonymy with Protothelenella corrosa.
KEYWORDS. – Floristics, forgotten species, herbaria, morphology, nomenclature, taxonomic
innovation.
INTRODUCTION
New species of lichens and lichenicolous fungi continue to be documented and described from
North America at a remarkable rate (Allen & Lendemer 2015, Allen & McMullin 2015, Brodo &
Lendemer 2015, Brodo & McCune 2017, Carlberg 2016, Cestaro et al. 2016, Cornejo et al. 2017, Driscoll
et al. 2016, Esslinger 2017, Esslinger et al. 2017, Fryday 2017, Gasparyan et al. 2017, Guzow-Krzemińska
et al. 2017, Holien et al. 2016, Knudsen et al. 2016, Lendemer 2016, Lendemer et al. 2016, Magain et al.
2016, McCune & Lumbsch 2017, McCune et al. 2018, Morse & Ladd 2016, Muggia et al. 2015,
Muscavitch et al. 2017, Myllys et al. 2016, Peterson & Goward 2016, Sanders & Lücking 2015, Seavey et
al. 2017, Tønsberg & Goward 2016). As a result, the number of taxa reported from North America north of
Mexico has steadily increased over time, and now stands at 5561 (Esslinger 2018). Although the discovery
of new and previously unreported taxa is a key component of biodiversity documentation, review and
revision of historical reports are similarly crucial components that nonetheless are often underappreciated
(see Lendemer 2015). In this context, there are many names currently included on the checklist of North
American lichens (Esslinger 2018) that were described or reported decades ago, in some cases even more
than a century ago, and yet have been little used, if at all, in modern times. As part of ongoing attempts to
address the status and taxonomic identity of such names, we present notes on four taxa described from
North America.
1
JAMES C. LENDEMER – Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY
10458-5126, U.S.A. – e-mail: jlendemer@nybg.org
2
RICHARD C. HARRIS – Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY
10458-5126, U.S.A. – e-mail: rharris@nybg.org
3
R. TROY MCMULLIN – Research and Collections, Canadian Museum of Nature, PO Box 3443 Stn “D”,
Ottawa, ON, K1P 6P4, Canada. – e-mail: tmcmullin@nature.ca

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MATERIALS AND METHODS
This study is based on specimens deposited in the herbaria of the New York Botanical Garden
(NY) and the Canadian Museum of Nature (CANL), supplemented by type specimens borrowed from UPS.
Georeferenced voucher data for all NY specimens examined can be accessed via the C.V. Starr Virtual
Herbarium at NY (http://sweetgum.nybg.org/science/vh/). Specimens were first studied dry using an
Olympus SZ-STB dissecting microscope. Microscopic morphology and anatomy were then studied using
an Olympus BX53 compound microscope and sections prepared by hand with a razor blade that were
mounted in water or Lugols solution (aqueous iodine). Chemistry was studied using standard chemical tests
(K, C, KC, P, UV) following Brodo et al. (2001) and supplemented by thin layer chromatography (TLC)
using solvents A and C following Culberson and Kristinsson (1970) but as modified for the peanut butter
jar by Lendemer (2011).
I. LECIDEA VIRGINIENSIS IS A SYNONYM OF BRYOBILIMBIA AHLESII
Bryobilimbia ahlesii (Körb.) Fryday, Printzen & S. Ekman, Lichenologist 46: 29. 2014. ≡ Biatora ahlesii
Korb., Parerg. Lichenol. p. 161. 1865. ≡ Lecidea ahlesii (Körb.) Nyl., Flora 55: 356. 1872. TYPE:
[GERMANY. BADEN-WÜRTTEMBERG.], Heidelberg, in sylvis “montanus” [montanis]
prope. 1852, W. Zwackh s.n. (H-NYL 20416[digital image!], neotype (designated by Meyer
2002)).
Syn. nov. = Lecidea virginiensis Calk. & Nyl., Bot. Gaz. 22: 333. 1896. TYPE: U.S.A. WEST
VIRGINIA. FAYETTE CO.: near Nuttallburg, 1893, on sandstone rock under the drip of a wet
cliff, L.W. Nuttall 1779 (H-NYL 20919!, lectotype designated here! [MBT-388347).
Discussion. – Lecidea virginiensis was first published as a nomen nudum in Millspaugh and
Nuttall (1896a: 181). The name was then validated in Millspaugh and Nuttall (1896b). Later, Millspaugh
(1913: 160) reproduced the original description with a reference to the earlier validation in the Botanical
Gazette. Importantly, all three references cited the same collection (Nuttall 1779) as the only material that
had been assigned to the name. Unfortunately, repeated attempts to locate L.W. Nuttall’s original material
that had clearly been assigned collection number 1779 were unsuccessful. It appears that Nuttall must have
sent material to W.W. Calkins, who then forwarded a specimen to W. Nylander for determination. The only
material of this species in H-NYL consists of a specimen sent by Calkins and marked “No. 168” by
Calkins, presumably referring to a numbered set of specimens that had been sent for identification to
Nylander. As has been documented elsewhere, the above scenario was common practice for several North
American workers (e.g., Calkins and J.W. Eckfeldt) who routinely sent collections made by others to
Nylander (LaGreca & Lumbsch 2001, Lendemer & Hewitt 2002). Thus, despite the fact that the specimen
in H-NYL lacks Nuttall’s collection number, we here select it as the lectotype because it was indisputably
studied by Nylander, described at length by Magnusson (1935), and comprises part of the original material
used to prepare the original description (Art. 9.4 in Turland et al. 2018).
Subsequent to its description, the name Lecidea virginiensis was occasionally used in the
published literature (e.g., Lowe 1939, Miller & Thomson 1959) and it remains on the North American
Checklist (Esslinger 2018). Despite this, the application of the name remains unclear and it does not appear
to have been used in recent times (i.e., since Miller & Thomson 1959). While preparing a checklist of the
lichens of Indiana, one of us (JCL) was spurred to determine whether L. virginiensis was a distinct and
overlooked species, or if it was a synonym of an older name that has become more widely used over time.
Examination of the type material at H-NYL (Fig. 1) revealed that L. virginiensis should be a treated as a
synonym of Bryobilimbia ahlesii.
Bryobilimbia ahlesii was reported from North America under the name L. ahlesii by Harris
(1995a) and has subsequently been found to be widespread in temperate eastern North America, including
the Appalachian Mountains (Fig. 2A). It has also been reported from Haida Gwaii in coastal British
Columbia (McCune 2017). The species is known to be an unusual member of Bryobilimbia in that it
typically occurs on wet or damp shaded sandstone (Arup 2004, Coppins & Fryday 2006, Fryday et al. 2014,
Purvis et al. 1992) and this perfectly matches the ecology that was originally described for L. virginiensis
(Millspaugh & Nuttall 1896a, b). The lectotype fits well with the published descriptions of L. ahlesii in
having a thin, continuous, esorediate thallus with small, reddish-brown to black, biatorine apothecia and

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Figure 1. Lecidea virginiensis (all from the lectotype). A and B, detail of the thallus and apothecia,
illustrating the variation in apothecial pigmentation. C, section of apothecium mounted in water. D, the
same apothecial section after application of KOH. Scales = 1.0 mm in A and B, 50 µm in C and D.
Porpidia-like asci. Nonetheless, it differs in having somewhat narrower ascospores (11.8–13.8 × 4.3–5.1
µm vs. 12–17(19) × (5.3)–6.4–7–(9) µm fide Coppins & Fryday 2006). Additionally, the apothecia entirely
lack the K+ green granules that have often been reported for the species. While these differences suggest
that further study of this group is needed, B. ahlesii is currently circumscribed to include forms that lack
K+ green granules in the apothecia (Coppins & Fryday 2006) and the slightly smaller size of the ascospores
may be attributable to the small number measured due to a desire to avoid damaging the lectotype. As such,
we here place L. virginiensis in synonymy with B. ahlesii. For additional heterotypic synonyms of B.
ahlesii (e.g., Lecidea delincta Nyl., L. valentior Nyl.), see Fryday et al. 2014).
Bryobilimbia sanguineoatra (Wulfen) Fryday, Printzen & S. Ekman is another species that is
morphologically similar to B. ahlesii and differs morphologically in its narrower ascospores. Although B.
sanguineoatra typically grows over mosses, it rarely also occurs on damp, siliceous rocks (Fryday et al.
2014). The width of the ascospores in B. sanguineoatra is comparable to that of Lecidea virginiensis (4.3–
5.1 µm in L. virginiensis vs. 3.0–4.5(–6.0) µm in B. sanguineoatra fide Fryday et al. 2014). Given the
ecology of the type of L. virginiensis we prefer to treat it as a synonym of B. ahlesii rather than as a
potential ecologically atypical form of B. sanguineoatra.
Selected specimens of Bryobilimbia ahlesii examined. – CANADA. NOVA SCOTIA.
COLCHESTER CO.: Economy River Wilderness Area, N end of Simpson Lake, 17.v.2004, on rock, W.R.
Buck 47211 (NY). QUEBEC. PONTIAC CO.: Gatineau Park, Church Hill, 22.v.2011, on rock, J.C.
Lendemer 28335 & C. Freebury (NY). U.S.A. ARKANSAS. FRANKLIN CO.: Ozark National Forest,
White Rock Wildlife Management Area, 8.vi.2000, on rock, W.R. Buck 37301 (NY). ILLINOIS.
JACKSON CO.: Shawnee National Forest, Pomona Natural Bridge, 1.x.1993, on rock, R.C. Harris 31294

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Figure 2. Geographic distributions of two species treated here based on specimens examined for this study.
A, distribution of Bryobilimbia ahlesii in eastern North America with the type locality of L. virginiensis in
black. B, distribution of Protothelenella corrosa in eastern North America with the type locality of
Thelenella humilis in black. Note the map for P. corrosa also includes literature occurrences published by
Fryday (2006, 2010).
(NY). MAINE. HANCOCK CO.: slopes of Lead Mountain, terminus of gravel road off ME9, 1.vi.2010, on
rock, J.C. Lendemer 23147 (NY). MARYLAND. CECIL CO.: S/E shore of Octoraro Creek, 0–1 mi S of
Porters Bridge, 26.v.2009, on rock, J.C. Lendemer 17813 (NY). NEW JERSEY. BERGEN CO.: Closter,
1876, on rock, C.F. Austin s.n. (NY). NEW YORK. GREENE CO.: Catskill Mountains, north end of Mink
Hollow and west slope of Sugarloaf Mountain, 8.x.2007, on rock, J.C. Lendemer 9796 (NY).
LIVINGSTON CO.: Town of Portage, Letchworth State Park, 26.ix.2015, on rock, J. Battaglia 2015-260
(NY). YATES CO.: Pen Yan, H.P. Sartwell s.n. (NY). NORTH CAROLINA. CLAY CO.: Nantahala
National Forest, Buck Creek Barren, 1–1.5 mi N of US 64 on Buck Creek Rd., 10.xi.2007, on rock, J.C.
Lendemer et al. 10820 (NY). SWAIN CO.: Great Smoky Mountains National Park, Juneywhank Falls, 14
.x.2010, on rock, E.A. Tripp et al. 1529 (NY); Great Smoky Mountains National Park, Beech Gap Trail
between Straight Fork Rd./Balsam Mountain Rd. and Hyatt Ridge Trail, 6.viii.2009, on rock, J.C.
Lendemer 19190 & E.A. Tripp (NY); Great Smoky Mountains National Park, Sunkota Ridge Trail between
Martins Gap and S spur trail to Indian Creek Trail, 21.vi.2011, on rock, J.C. Lendemer 29445 & N.
Davoodian (NY); Great Smoky Mountains National Park, 0.25 mi N of terminus of Bone Valley Trail,
3.v.2015, on rock, J.C. Lendemer 44651 & E.A. Tripp (NY). RHODE ISLAND. PROVIDENCE CO.:
Town of Lincoln, Lime Rock Nature Preserve, 17.ix.2006, on rock, W.R. Buck 51105 (NY).
PENNSYLVANIA. BRADFORD CO.: State Game Lands No. 36, Falls Creek, 1 mi N of confluence with
Schrader Creek, 18.v.2009, on rock, J.C. Lendemer 17292 & R.C. Harris (NY). CAMERON CO.: Elk
State Forest, Fourmile Run Rd., 3.25 mi N of jct w/ PA 155, 2.ix.2010, on rock, J.C. Lendemer 24432
(NY). LUZERNE CO.: Rickets Glenn State Park, northeast of Red Rock, The Glens Natural Area, along
Kitchen Creek, 17.iv.2004, on rock, J.C. Lendemer 2366 & J.A. Macklin (NY). LYCOMING CO.: State
Game Lands No. 133, E shore of Lycoming Creek, N-facing slopes of Batys Mountain between Debois
Hollow and Shoemaker Run, 12.v.2009, on rock, J.C. Lendemer 16649 & D. Atha (NY). WAYNE CO.:
Dripping Cliffs TNC Preserve, 2.vii.2008, on rock, J.C. Lendemer 12372 (NY). WYOMING CO.: State
Game Lands No. 57, Dutch Mountain, 21.vii.2008, on rock, J.C. Lendemer 13692 (NY). YORK CO.: W-
shore of Susquehanna River, 0.5 mi S of jct of River Rd. & PA 372, 28.v.2009, on rock, J.C. Lendemer
17977 (NY). TENNESSEE. COFFEE CO.: Old Stone Fort State Archeological Area, vi.1988, on rock,
J.P. Dey 17588 (NY). SEVIER CO.: Great Smoky Mountains National Park, Chimney Tops Trail,
28.vi.2010, on rock, W.R. Buck 56273 (NY). VIRGINIA. GRAYSON CO.: Jefferson National Forest, Mt.
Rogers National Recreation Area, Comers Creek, 5.iv.2008, on rock, G.B. Perlmutter 1283 (NY).

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Figure 3. Holotypes of Lecidea names based on specimens from Great Smoky Mountains National Park. A
and B, external morphology of L. deminutula. C and D, external morphology of Lecidea olivacea var.
inspersa. Scales = 2.0 mm in C, 1.0 mm in A and D, 0.5 mm in B.
II. LECIDEA DEMINUTULA IS A SYNONYM OF LECIDELLA ENTEROLEUCELLA
Lecidella enteroleucella (Nyl.) Hertel, Khumbu Himal 6: 330. 1977. ≡ Lecidea enteroleucella Nyl. in
Nylander & Crombie, J. Linn. Soc., Bot. 20: 67. 1883. TYPE: JAPAN. NAGASAKI PREF.:
Nagasaki, A.C. Maingay s.n. (H-NYL 15625 [digital image!], holotype).
Syn. nov. Lecidea deminutula H. Magn., in Degelius, Ark. f. Bot. 30A(3): 35. 1942. TYPE: U.S.A.
TENNESSEE. SEVIER CO.: Great Smoky Mountains National Park, near Laurel Falls, 760 m.,
16.ix.1939, on rock, G. Degelius s.n. (UPS [L-74933]!, holotype).
Discussion. – Lecidea deminutula was described from the southern Appalachian Mountains of
eastern Tennessee based on material collected on non-calcareous rocks (Degelius 1942). The name does not
appear to have been used subsequently in North American literature, and it remains on the North American
Checklist (Esslinger 2018). Although a placement in Lecidella would have been suggested based on the
discussion presented by Magnusson (in Degelius 1942), the name does not appear to have been treated in
any of the primary works dealing with the genus (e.g., Andreev & Baibulatova 1986, Andreev et al. 2003;
Inoue 1982, 1997, 2000; Kantvilas & Elix 2013; Knoph 1990; Knoph & Leuckert 1994, 1997, 2000, 2004;
Leuckert et al. 1990, 1992; Rambold 1989).
We examined the holotype at UPS (Fig. 3A–B) and found that it corresponds well to the current
concept of Lecidella enteroleucella (Knoph & Leuckert 1994, 2004), a species that is widespread in
temperate eastern North America (Knoph & Leuckert 1994; Fig. 4B). The holotype has a thin, areolate,

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Figure 4. Geographic distributions of two species treated here based on specimens examined for this study.
A, distribution of Lecidella elaeochroma in southeastern North America with the type locality of Lecidea
olivacea var. inspersa in black. B, distribution of Lecidella enteroleucella in eastern North America with
the type locality of Lecidea deminutula in black.
esorediate thallus, hyaline hypothecium, hyaline hymenium that is not inspersed with oil droplets, a dark
olive-brown epihymenium, and ascospores that measure 9.4–12.2 × (3.7)–4.4–6.1 µm in size. Thin layer
chromatography of the holotype detected atranorin together with two xanthones, likely arthothelin and
thuringione. The identities of the latter two substances were not subsequently confirmed with standards as
the type specimen comprises a single rock fragment and we wanted to avoid repeated sampling of the
limited available material. Nonetheless, given the morphological characters present in the type, the
occurrence on non-calcareous rocks, and the presence of both atranorin and xanthones, the assignment to L.
enteroleucella seems clear. As such we formally place the names in synonymy here.
Lecidella enteroleucella has been little discussed in the published literature subsequent to Knoph
and Leuckert (1994). Harris and Ladd (2005) included it in an account of Ozark lichens and compared the
species to L. stigmatea (Ach.) Hertel & Leuckert, distinguishing the species by the degree of immersion of
the apothecia in the thallus (immersed in L. enteroleucella vs. sessile in L. stigmatea). Based on the
specimens examined for this study, the apothecia of L. enteroleucella can vary from immersed to sessile,
the latter exemplified by the type of L. deminutula. Lecidea stigmatea differs morphologically in having
larger ascospores (11–17 × 6–9 µm, fide Knoph & Leuckert 2004) and chemically from L. enteroleucella in
producing zeorin as an accessory to atranorin or lichexanthone (Knoph & Leuckert 2004).
Selected additional specimens of Lecidella enteroleucella examined (all from non-calcareous
rocks). – U.S.A. ALABAMA. DEKALB CO.: Little River Canyon National Preserve, Lower Two Mile
Trail from scenic highway to river, 22.x.2012, E.A. Tripp 4004 (NY). JACKSON CO.: Bucks Pocket State
Park, S facing slopes above N shore of South Sauty Creek, 15.xii.2016, E. Tripp 6087 & J.C. Lendemer
(NY). ARKANSAS. FRANKLIN CO.: Ozark National Forest, summit of White Rock Mountain,
24.iv.1988, R.C. Harris 21429 (NY). PERRY CO.: Ouachita National Forest, vicinity of Goat Bluff along
north side of South Fourche LaFave River, 6.x.2010, J.C. Lendemer et al. 26041 (NY). ILLINOIS.
RANDOLPH CO.: Piney Creek Ravine Nature Preserve, 7.xi.2004, R.C. Harris 50563 (NY). KANSAS.
CHEROKEE CO.: Schermerhorn Park, just S of Galena on KS 26, 21.x.2000, R.C. Harris 44390 (NY).
MONTGOMERY CO.: 0.5 mi N, 5.5 mi W Liberty, S side of Montgomery Co. State Park, 8.xi.2006, C.A.
Morse 14375 (NY). MISSOURI. REYNOLDS CO.: Johnson Shut-Ins State Park, along East Fork Black
River E of CR-N, 9.x.1997, M.S. Cole 7457 (NY). WAYNE CO.: Sam A. Baker State Park, Mudlick
Mountain, 15.x.2003, R.C. Harris 48379 (NY). NEW JERSEY. MERCER CO.: Hopewell Township,
Belle Mountain adj to NJ-29 & Delaware River, 10.iii.2016, D.P. Waters 1800 (NY). NORTH
CAROLINA. CLAY CO.: Nantahala National Forest, stream bed/valley of Buck Creek, 1–1.5 mi N of US
64 on Buck Creek Rd., 10.xi.2007, J.C. Lendemer et al. 10458 (NY). MONTGOMERY CO.: Uwharrie
National Forest, Barnes Creek Bluffs, N of Dark Mountain, 19.iv.2008, G.B. Perlmutter et al. 1399 (NY).

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OKLAHOMA. ADAIR CO.: just NE of Stilwell City Lake, 1.xi.2000, R.C. Harris 44512 (NY).
VIRGINIA. ALBEMARLE CO.: Shenandoah National Park, South District, Dundo Picnic Area,
9.ix.2016, Z. Muscavitch 206 & J.C. Lendemer (NY). WEST VIRGINIA. GREENBRIER CO.: Wadea
Draft, NE of CR 15-2, 1.x.2000, R.C. Harris 44037B (NY). POCAHONTAS CO.: Monongahela National
Forest, upper slopes of Spruce Mountain, vicinity of Blue Knob, 20.x.2007, J.C. Lendemer 9920 & A.
Moroz (NY).
III: LECIDEA OLIVACEA VAR. INSPERSA IS A SYNONYM OF LECIDELLA ELAEOCHROMA
Lecidella elaeochroma (Ach.) M. Choisy, Bull. mens. Soc. linn. Soc. Bot. Lyon 19: 19. 1950. ≡ Lecidea
parasema var. elaeochroma Ach., Methodus, Sectio prior (Stockholmiæ): 36. 1803. ≡ Lichen
myriocarpus * elaeochroma (Ach.) Lam., Encycl. Méth. Bot., Suppl. 3: 371. 1813. ≡ Lecidea
elaeochroma (Ach.) Ach., Syn. meth. lich. (Lund): 18. 1814. ≡ Lecidea parasema var. enteroleuca
(Ach.) Nyl., Act. Soc. linn. Bordeaux 21: 370. 1856. ≡ Biatora olivacea var. elaeochroma (Ach.)
Hepp, Flecht. Europ.: no. 247. 1857. ≡ Lecidea parasema ß elaeochroma (Ach.) Harm., Bull.
Séanc. Soc. Sci. Nancy, Sér. 2 33: 77. 1899 ≡ Lecidea parasema subf. elaeochroma (Ach.) Harm.,
Bull. Séanc. Soc. Sci. Nancy, Sér. 2 33: 76. 1899 ≡ Lecidea olivacea f. elaeochroma (Ach.)
Mereschk., Annuaire Conser. et Jard. bot. Genève 21: 209. 1919. TYPE: see below.
Syn. nov. = Lecidea olivacea var. inspersa Degel., Ark. f. Bot. 30A(3): 39. 1942. TYPE: U.S.A.
TENNESSEE. SEVIER CO.: Great Smoky Mountains National Park, near The Chimneys, 850
m., 19.ix.1939, on Celtis, G. Degelius s.n. (UPS [L-587982]!, holotype).
Discussion. – Lecidea olivacea var. inspersa was described by Degelius (1942) based on
corticolous material collected in the southern Appalachian Mountains of eastern North America. As was the
case for L. deminutula, the protologue suggests a placement in Lecidella. Nonetheless, the name does not
appear to have been treated in any of the primary works on that genus (e.g., Andreev & Baibulatova 1986,
Andreev et al. 2003; Inoue 1982, 1997, 2000; Kantvilas & Elix 2013; Knoph 1990; Knoph & Leuckert
1994, 1997, 2000, 2004; Leuckert et al. 1990, 1992; Rambold 1989). Likewise, following its description
the name does not appear to have been used again in the North American literature.
We examined the holotype deposited in UPS (Fig. 3C–D) and found that it is a species of
Lecidella that morphologically and chemically agrees well with current delimitations of L. elaeochroma
(e.g., Knoph & Leuckert 1997, 2004), a species that occurs sporadically throughout the southern
Appalachian Mountains (Fig. 4A). Specifically, the holotype has a thin, esorediate thallus, dark blue-black
apothecia with a blue-green epihymenium, hyaline hymenium densely inspersed with oil droplets, yellow-
brown pigmented hypothecium and exciple, and ascospores that measure 12.7–15.6 × 7.0–9.6 µm. The
chemistry of the holotype was studied with both spot tests and thin layer chromatography, using the former
we found the thallus was K+ yellow, C-, KC-, P- and UV-, while using the latter, we detected atranorin and
an unidentified xanthone. Due to the small size of the holotype we did not subject the material to further
chemical analyses, and while such studies should be carried out to identify the xanthone, we suggest that
methods such as HPLC should be used as they require minimal material. Following Knoph and Leuckert
(2004), L. elaeochroma is morphologically nearly identical to L. euphorea (Flörke) Hertel, differing in the
xanthones it produces. Lecidella elaeochroma has widely been reported to react C+ orange, albeit often
weakly so, while L. euphorea has been reported to be C- (e.g., Purvis et al. 1992). However the differences
between the species do not appear to be clear in North America (Knoph & Leuckert 2004) and further study
of species delimitation in this group is clearly needed. We here treat Lecidea olivacea var. inspersa as a
synonym of L. elaeochroma, recognizing that there is a chance it could prove to be better accommodated as
a synonym of L. euphorea when the group is studied in more detail in the future.
It should be noted that after extensive review of the literature we were unable to locate a published
typification for Lecidella elaeochroma. This is despite the fact that the species is considered to be
widespread and abundant, especially in Europe (e.g., Purvis et al. 1992). There are three sheets with
material identified as L. elaeochroma in Acharius’ herbarium (H-ACH 140A-F, H-ACH 141, H-ACH 154).
Of these, H-ACH140A was annotated by Leuckert and colleagues as containing diploicin and
lichexanthone. Given that there are multiple specimens comprising original material which have not been
chemically analyzed, and that the one specimen annotated with chemical data does not agree with

97
published chemical accounts of the species (e.g., Knoph & Leuckert 2004), we defer lectotypification until
the material can be studied in detail.
Selected additional specimens of Lecidella elaeochroma examined. – GERMANY.
RHEINLAND-PFALZ. Kr. Cochem. Naturschutzgebiet Dortebachtal, along trail from parking area to
waterfall, ca. 1 km E of village of Klotten along Hwy 49, 30.i.2010, on Acer, W.R. Buck 55939 (NY).
U.S.A. NORTH CAROLINA. HAYWOOD CO.: Great Smoky Mountains National Park, E slopes above
(E of) Caldwell Fork, along E portions of Boogerman Loop Trail, 4.viii.2009, on Liriodendron, E. Tripp
575 & J.C. Lendemer (NY); Pisgah National Forest, Shining Rock Wilderness, middle E-facing slopes of
Old Butt Knob, 7.vi.2015, on Acer, J.C. Lendemer 45338 & J.L. Allen (NY). MACON CO.: Nantahala
National Forest, NW slope of Cole Mountain, 6.xii.2016, on dead Quercus, J.C. Lendemer 48616 (NY).
SWAIN CO.: Great Smoky Mountains National Park, Beech Gap Trail between Straight Fork Rd./Balsam
Mountain Rd. and Hyatt Ridge Trail, 6.viii.2009, on Acer, J.C. Lendemer 19240 & E. Tripp (NY).
TENNESSEE. BLOUNT CO.: Great Smoky Mountains National Park, between Flint Gap and Hannah
Mountain on Hannah Mountain Trail, 9.viii.2012, on Acer, E. Tripp 3587 & J.C. Lendemer (NY). COCKE
CO.: Great Smoky Mountains National Park, Gabes Mountain Trail, 0–2 mi E of jct with Maddron Bald
Trail, 5.viii.2009, on fallen branch, E. Tripp 655 & J.C. Lendemer (NY).
IV: THELENELLA HUMILIS IS A SYNONYM OF PROTOTHELENELLA CORROSA
Protothelenella corrosa (Körb.) H. Mayrhofer & Poelt, Herzogia 7: 42. 1985. ≡ Limboria corrosa Körb.,
Syst. Lich. Germ., p. 376. 1855. ≡ Acrorixis corrosa (Körb.) Trevis., Conspect. Verruc., p. 15.
1860. ≡ Polyblastia corrosa (Körb.) Arnold, Flora 53: 19. 1870. ≡ Microglaena corrosa (Körb.)
Arnold, Verh. zool.-bot. Ges. Wien 27: 555. 1877. ≡ Verrucaria corrosa (Körb.) Vain.,
Meddeland. Soc. Fuana Fl. Fenn. 10: 182. 1883. ≡ Thelenella corrosa (Körb.) Vain., Acta Soc.
Fauna Fl. Fenn. 49(2): 353. 1921. ≡ Microglaena nericiensis var. corrosa (Arnold) Hellb.,
Nerekes Lafflora, p. 123. 1871. TYPE (fide Mayhofer & Poelt 1985: 42): “Polen: An versteckten
Granitblöcken im Melzergrunde im Riesengebirge, 1841, W.G. Korber” (L[n.v.], missing).
Syn. nov. = Thelenella humilis R.C. Harris, More Florida Lichens, p. 165. 1995. U.S.A. NEW YORK.
WARREN CO.: Crane Mountain, hardwoods below to spruce woods near summit, 18 Sept. 1993,
on boulder in trail, R.C. Harris 30577 (NY!, holotype).
Discussion. – Thelenella humilis was described from a single location in the Adirondack
Mountains of New York by Harris (1995b). In the more than two decades since its description no additional
specimens have been referred to as this taxon and it has rarely been mentioned in the literature (e.g., Morse
2016). Given that this species was described from northeastern North America and only located once, the
authors have been curious as to whether it might represent a rare species or a crustose lichen that has
otherwise been overlooked in the field. During a visit to New York in 2015 one of us (TM) examined the
type of T. humilis and brought it to the attention of the other authors. We quickly realized that the specimen
corresponds to typical material of Protothelenella corrosa (Fig. 5). Indeed, P. corrosa is one of the few
peritheciate crustose lichens in eastern North America that occurs on non-calcareous rocks and has hyaline,
muriform ascospores and octosporous asci. Comparison between the original description of T. humilis and
published accounts of P. corrosa (e.g., Mayrhofer & Poelt 1985, Lendemer et al. 2009) further confirmed
the identification based on the presence of submuriform ascospores and a C+ pink-red thallus. As is
evidenced by the comparative specimens cited below, P. corrosa is widely distributed in northeastern
North America and the type of T. humilis was collected well within the documented range of P. corrosa
(Fig. 2B). We here place T. humilis in synonymy with P. corrosa.
Selected specimens of Protothelenella corrosa examined. – CANADA. NEW BRUNSWICK.
PENNFIELD PARISH: W of Jake Lee Rd., W and S of small unnamed lake at head of unnamed tributary
of Love Lake Brook, 1.v.2011, on rock, W.R. Buck 57703 (NY). NEWFOUNDLAND AND
LABRADOR. NEWFOUNDLAND: Avalon Peninsula, Butterpot Provincial Park, E side of Big Otter
Pond, 9.ix.2007, on rock, J.C. Lendemer 10162 (NY). NOVA SCOTIA. COLCHESTER CO.: Economy
River Wilderness Area, vicinity of Economy Falls, 14.v.2004, on rock, W.R. Buck 47036 (NY). QUEBEC.
M.R.C. DE LA HAUTE-GASPÉSIE: Parc national de la Gaspésie, Sentier du Mont Jacques-Cartier, from

98
Figure 5. Morphological comparison of Thelenella humilis (A,C,E) (all from the holotype) with
Protothelenella corrosa (B,D,F) (all from Lendemer 32525). A and B, gross morphology of the thallus. C-
F, detail of the ascomata. Scales = 1.0 mm in A-D, 0.5 mm in E and F.

99
trailhead to Lac à René, 3.vi.2012, on rock, W.R. Buck 59631 (NY); Mont Albert, c. 1 km W of the park
office, along trail to the N summit, 4.vii.2012, on rock, R.T. McMullin 11969 (CANL). U.S.A.
MASSACHUSETTS. BERKSHIRE CO.: Mount Washington, Mount Everett State Forest, Guilder Pond
Loop, 8.ix.2012, on rock, E. Lay 12-0117 (NY). MAINE. HANCOCK CO.: Donnell Pond Maine Public
Reserve Lands, SE slopes of Schoodic Mountain, 7.vi.2012, on rock, J.C. Lendemer 32430 & A. Moroz
(NY). PENNSYLVANIA. HUNTINGDON CO.: Rothrock State Forest, Colerain Picnic Area, 21.iv.2008,
on rock, W.R. Buck 53522 (NY).
ACKNOWLEDGEMENTS
The study is a result of NSF DOB Awards 1542639 (to PI-Lendemer) and 1542629 (to PIs-Kane, McCain
and Tripp). We thank the curators of UPS for the loan of specimens used in this study. Leena Mylyys and Teuvo Ahti
provided helpful discussion of the type of Lecidella elaeochroma. Two reviewers provided helpful comments and
revisions to the manuscript.
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