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Opuscula Philolichenum, 15: 22-36. 2016.
*pdf effectively published online 21July2016 via (http://sweetgum.nybg.org/philolichenum/)
Two new species of Thelenella and new reports from the
Great Plains of central North America, with a worldwide
key to the genus
CALEB A. MORSE
1
ABSTRACT. – Two new species of Thelenella are described from central North America.
Thelenella calcicola occurs on calcareous rocks in Kansas and Missouri, and produces irregularly
submuriform, 7–11 × 1-septate, colorless ascospores. Thelenella nubifera, based on specimens previously
referred to T. luridella, occurs on sandstone in Georgia, Kansas, Missouri, and Oklahoma, and produces
muriform, 6–9 × 3–4-septate, grey or brown pigmented ascospores. A third species from northwestern
South Dakota is characterized but not formally described. Thelenella luridella is tentatively excluded from
North America. Thelenella brasiliensis is reported new for Kansas, southwest Oklahoma, and Texas; T.
modesta is reported new for Kansas and North Dakota; T. muscorum var. muscorum is reported new for or
Kansas and Oklahoma; and T. pertusariella is reported new for Kansas. A worldwide key to Thelenella is
provided.
KEYWORDS. – Amyloidy, Altamaha Formation, Cross Timbers, biogeography, ecoregions,
Ozarks, taxonomy.
INTRODUCTION
Thelenella Nyl. (Lecanoromycetes: Ostropomycetidae: Thelenellaceae) comprises a genus of
crustose lichens with a chlorococcoid photobiont, perithecia that are often immersed and in most species
lack an involucrellum, a hamathecium of slender, branched and anastomosing paraphyses and simple or
branched periphysoids, thick-walled, bitunicate asci with a more or less well-developed, KI‒ ocular
chamber, 8 or fewer, mostly colorless, submuriform to muriform ascospores, and filiform conidiospores
(Mayrhofer 1987, Mayrhofer & McCarthy 1991, Fryday & Coppins 2004). The genus includes 35
corticolous, foliicolous, muscicolous, and saxicolous taxa (Mayrhofer 1987, Mayrhofer & McCarthy 1991,
Harris 1995, Kalb 1995, Etayo & Mayrhofer 2003, Fryday & Coppins 2004, Pinokiyo & Singh 2006,
Aptroot et al. 2015), although the inclusion of some taxa has been, or continues to be, contentious. One
species, initially described as a non-lichenized Thelenella (Aptroot 1999), has subsequently been shown to
belong to a monotypic genus in the Trypetheliaceae, Aptrootia Lücking & Sipman (Lücking et al. 2007).
Recent inclusion of three polysporous species by Aptroot and Schumm (2012) has been rejected by
Knudsen and Kocourková (2013). Transfer of several species of the mostly foliicolous genus
Aspidothelium Vainio to Thelenella by Harris (1995) and Farkas and Sipman (1997) was disputed by
Lücking (2008), who postulated a distant relationship between Aspidothelium and the remainder of the
Thelenellaceae. However, molecular sequence data have not yet been generated for Aspidothelium, and the
placement of this genus remains uncertain. Aspidotheliaceae is presently included as a family incertae sedis
of Ascomycota by Lumbsch and Huhndorf (2009). Thelenella is represented in North America by 18
species (Esslinger 2015), including two species of Aspidothelium.
Saxicolous members of the genus typically occur on non-calcareous rocks, and most are limited to
maritime or oceanic climates (Mayrhofer & McCarthy 1991), so it was surprising to find a species growing
on the sheltered face of limestone outcrops along the eastern edge of the Great Plains. In reviewing other
members of the genus as part of preparing a description of this taxon, it became evident that a second
species found in southeastern North America, which produces distinctive, grey to brownish ascospores,
1
CALEB A. MORSE – Biodiversity Institute, R.L. McGregor Herbarium, University of Kansas, 2045
Constant Ave., Lawrence, KS 66047, U.S.A. – e-mail: cmorse@ku.edu
23
could not be placed satisfactorily in any known Thelenella. Both species are described here as new to
science. A third species, known from a single small specimen from South Dakota, is discussed but not
formally described. In addition, several species are newly reported for Great Plains states in North
America. Further, because Thelenella has grown considerably since Mayrhofer’s (1987) revision, a
worldwide key to the genus is provided.
MATERIALS AND METHODS
Chemical analysis was conducted using standard spot tests (reagents are abbreviated following
Brodo et al. (2001)) and Thin Layer Chromatography (TLC). TLC was carried out at KANU using solvent
systems A and C following the methods of Orange et al. (2001). Morphological study was carried out on
hand sections prepared with a razor blade and mounted in water. Medullary cells were stained with 0.3%
Lugols iodine (I) after pretreatment with 10% potassium hydroxide (K). Other microscopic characters were
observed in water and images were captured with a Nikon Eclipse 80i microscope outfitted with a
Lumenera INFINITY-32 digital camera and measured to the nearest 0.1 μm with Lumenera INFINITY
ANALYZE imagining software. Measurements are presented as a simple range or, where sufficient
material allowed, as the average (xˉ) +/- one standard deviation (SD), bounded by the smallest and largest
observed values, and followed by the sample size (n) (i.e., (smallest observed) xˉ-1SD–xˉ–xˉ+1SD (largest
observed) [n]). Taxonomic authorities are not included for the names of associated species as these can
readily be obtained from Esslinger (2015).
TAXONOMIC SECTION
Thelenella calcicola C.A. Morse sp. nov.
MYCOBANK #817440. FIGURES 1A-E.
DIAGNOSIS. – Similar to Thelenella pertusariella (Nyl.) Vainio, but saxicolous on calcareous
rocks, with a thin, continuous thallus, and irregularly submuriform, (5) 7–11 (12) ×1 (–2)-septate
ascospores, (18.5–)20.5–24.5–28.5(–33) × (5.4–)6.3–7.3–8.3(–9.8) μm.
TYPE: U.S.A. KANSAS. JEFFERSON CO.: 0.7 mi S, 7.2 mi E of Williamstown, University of
Kansas Ecological Reserves: Nelson Environmental Study Area and Rockefeller Experimental Tract: unit
4018, near 39.05132°N 95.19765°W, elev. 1000–1050 ft., N and NW-facing limestone outcrops and low
cliffs in brushy, mesic, mixed hardwood forest on moderate slopes along WNW-trending ravine at head of
unnamed tributary to Mud Creek, soil rocky, with some chert residuum along top of slope, on N-facing
outcrops and boulders and base of W-facing outcrops, 18.i.2016, C.A. Morse 24956 (KANU 399092!,
holotype; NY!, isotype).
DESCRIPTION. – Thallus epilithic, thin, continuous, membranous to cracked-areolate, matte or
somewhat lustrous, finely mottled greenish grey and white or tan to yellowish brown, surface smooth,
frequently dimpled with collapsed or eroded, greenish or brownish pigmented ascomata; medulla K/I–.
Photobiont chlorococcoid, the cells (7–)9–12(–15) μm in diameter. Ascomata dispersed or occasionally in
groups of 2–3, half-immersed in the thallus or immersed in hemispherical verrucae; verrucae 0.2–0.4 mm
in diameter, concolorous with thallus except for brown ostiole, densely inspersed with colorless to
brownish, ± rectilinear crystals (these POL+ yellowish, persisting in K); involucrellum lacking. Centrum
0.15-0.3 mm in diameter, exciple ca. 20 μm thick laterally, hyaline below, pale (reddish to greenish) brown
above (dull brown in K); periphysoids simple to sparsely branching, ca. 30–36 × 1.5 μm; paraphyses thin,
ca. 1 μm wide, abundantly branching. Asci narrowly pyriform to cylindrical, 50–85 × 11–16 μm, with (4?)
8 ascospores, KI–, with a distinct ocular chamber. Ascospores narrowly ellipsoid, spindle-shaped, or
typically clavate, with the proximal (and occasionally, distal) 2–4 cells noticeably narrowed and often with
pointed ends, colorless, irregularly submuriform, with (5–)7–11(–12) transverse septa and the median 1–5
cells and often the distal cell with 1 longitudinal septum each, or rarely with one median cell with 2
longitudinal septa, (18.5–)20.5–24.5–28.5(–33) × (5.4–)6.3–7.3–8.3(–9.8) μm, L/W ratio = (2.4–)2.9–3.4–
3.9(–4.6) [n = 53], immature ascospores occasionally with a perispore ca. 1–1.5 μm thick. Pycnidia black,
ca. 0.5 mm in diameter, upper wall with dark greyish green pigment (dull brown in K); conidia filiform,
gently curved to arcuate, ca. 14–19 × < 1 μm.
24
25
CHEMISTRY. – Spot tests of thallus with K, C, P all negative; thallus UV-. No substances detected
by TLC.
ETYMOLOGY. – The epithet calcicola is Latin (from calci- limestone + colo to inhabit) and refers
to the preferred substrate of this species.
ECOLOGY AND DISTRIBUTION. – Thelenella calcicola is known from a handful of sites in the
Central Tallgrass Prairie and Osage Plains/Flint Hills Prairie ecoregions of eastern Kansas and western
Missouri (TNC 2007; figure 2A), where it occurs on sheltered or shaded north, east, and west-facing
limestone outcrops and on larger limestone boulders in mesic, mixed oak-hickory-basswood forest from
850 to 1050 feet (259–320 meters) above sea level. Associated species include Bacidia granosa, B. suffusa,
Bacidina egenula, Bagliettoa baldensis, Botryolepraria lesdainii, Caloplaca sp. of Harris and Ladd (2005),
Dermatocarpon muhlenbergii, Endocarpon diffractellum, Gyalecta jenensis, Gyalolechia flavovirescens,
Lecania perproxima auct., Lepraria finkii, Squamulea subsoluta, Verrucaria calkinsiana, and V. alutacea.
Given its predilection for mesic habitats, it seems unlikely that T. calcicola is restricted the eastern edge of
the Great Plains. Its mottled thallus gives the species the aspect of the deteriorated thalli of Bacidia granosa
or Bagliettoa baldensis in the field, making it easy to overlook; additional populations should be sought in
appropriate habitats elsewhere in eastern North America.
DISCUSSION. – Following Mayrhofer and Poelt (1985) and Mayrhofer (1987), the ascus type of
both of the species described here might best be described as Chromatochlamys-type, with asci typically
showing a pronounced indentation of the endoascus at the tholus, at least during some stages of
development. However, whether one sees the character diagnostic for Chromatochlamys or Thelenella-type
asci appears to depend on the developmental stage of the ascus being observed. This agrees with earlier
findings by Harris (1995), Fryday and Coppins (2004), and Schmitt et al. (2005), who pointed out that asci
in members of Thelenella—including the type species T. modesta (Nyl.) Nyl.—frequently show a distinct
ocular chamber, and that Chromatochlamys and Thelenella-type asci may not be discrete types.
The variably submuriform ascospores of Thelenella calcicola are reminiscent of those found in T.
pertusariella, a corticolous species that often has a whitish, fleck-like thallus and tan, partially immersed
ascomata (Mayrhofer 1987). Thelenella pertusariella is rare in the eastern Great Plains, documented from
few collections on Quercus (Harris & Ladd 2005, and discussed below). Several other taxa produce
ascospores similar to those of T. calcicola. Thelenella sychnogonioides (Zahlbr.) R. C. Harris, a corticolous
species known from coastal California and New South Wales, differs in producing ascospores that are
broader than those of T. calcicola (20–30 × 7–11 μm, as T. harrisii H. Mayrhofer in Mayrhofer 1987), with
1–2 longitudinal septa. Another corticolous species, T. justii (Servit) H. Mayrhofer & Poelt, known from
Greece, produces ascospores that are much longer than those of T. calcicola (35–50 × 9–12 μm fide
Mayrhofer & Poelt 1985) and also have 1–2 longitudinal septa. Thelenella larbalestieri (A.L. Sm.) Coppins
& Fryday, a species on siliceous rocks known only from the United Kingdom, has a brownish thallus and
ascospores that are also much longer than those of T. calcicola (44–60 × 10–14 µm fide Mayrhofer & Poelt
1985; see also Fryday & Coppins 2004), and transversely septate to submuriform, with 0–2 longitudinal
septa. Thelenella vezdae (H. Mayrhofer & Poelt) Coppins & Friday, presently known only from Austria,
differs in being primarily muscicolous, or rarely corticolous on dead conifers (Mayrhofer & Poelt 1985,
Fryday & Coppins 2004), as well as in having ascospores that are slightly broader than those of T. calcicola
(22–30 × 7–10 µm fide Mayrhofer & Poelt 1985), with 0–2 longitudinal septa.
Figure 1 (Page 24). A–E, Thelenella calcicola. A, Thallus (Morse 24956 [holotype], KANU; arrows point
to ascomata), B, mature spores in ascus, after staining with K/I (Morse 24881, KANU), C, ascospores, with
halo (Morse 24956 [holotype], KANU; arrow points to distended area of halo), D, ascospores (Morse
24951, KANU), E, conidia (Morse 24961, KANU). F–J, Thelenella nubifera. F, specimen with tan thallus
(Morse 25135, KANU [holotype]; arrows point to ascomata), G, specimen with verrucose, greenish grey
thallus (Morse 24439, KANU), H, section through confluent ascomata, showing mature (grey) ascospores
and overmature (brown) ascospores (Morse 24439, KANU), I, mature ascospores with 2 longintudinal
septa (Morse 23280, KANU), J, mature ascospores with 3 longitudinal septa (Buck 48651, NY). Scales in
A, F, and G = 1 mm; scales in B, C, D, E, I and J = 20 μm; scale in H = 100 μm.
26
Among the saxicolous taxa documented from central North America, Thelenella calcicola is
unique in having narrow, irregularly submuriform ascospores and in occurring on strongly calcareous
substrates. Thelenella brasiliensis (Müll. Arg.) Vainio, which is known from a handful of Great Plains
collections on non-calcareous rocks, produces muriform ascospores with 2–3 longitudinal septa, which are
also broader than those of T. calcicola (9–13 μm wide fide Mayrhofer 1987). Thelenella nubifera
(described and discussed below) differs in having larger, muriform, brown or gray spores, and in occurring
on sandstone. Thelenella sastreana R.C. Harris, known from a few specimens collected in Puerto Rico and
Louisiana, produces ascospores that are slightly broader than those of T. calcicola (8–11 μm fide Harris
1995), with 1–2 longitudinal septa. In addition to producing slightly broader, muriform ascospores, T.
sastreana is reported to have larger asci (85–130 × 15–20 μm), and occurs on sandstone (Harris 1995).
Additional specimens examined. – U.S.A. KANSAS. DOUGLAS CO.: ca. 0.5 mi N, 5.6 mi W of
Lecompton, along S side of Scenic River Rd (= N 2190 Rd), 39.05°N 95.50°W, 28.iv.2015, C.A. Morse et
al. 24489 (KANU). JEFFERSON CO.: ca. 1 mi S, 3 mi E Ozawkie, E side of Perry Lake, along Old
Military Trail, near 39.21°N 95.41°W, 11.x.2015, C.A. Morse et al. 24866 (KANU, hb. Ladd, NY).
JOHNSON CO.: ca. 1.5 mi N, 1 mi W of Aubry, Overland Park Arboretum and Botanical Garden, 38.80°N
94.69°W, 2.xi.2015, C.A. Morse 24881 (KANU, NY). MIAMI CO.: 0.25 mi N, 2 mi E of Jingo, North La
Cygne State Fishing Lake and Wildlife Area, along W side of Rockville Rd, N of intersection Rockville Rd
and 399th St, 37.41°N 95.66°W, 11.xii.2015, C.A. Morse 24951 (KANU). MISSOURI. JACKSON CO.:
Kansas City, Blue River Glades Natural Area, along E side of Blue River Rd (Blue River Co Pkwy), ca.
0.75 road miles N of intersection of Blue River Rd and E 87th St, 38.98°N 94.54°–94.53°W, 29.i.2016,
C.A. Morse 24961 (KANU).
Thelenella nubifera C.A. Morse sp. nov.
MYCOBANK #817441. FIGURES 1F-G.
DIAGNOSIS. – Similar to Thelenella luridella (Nyl.) Mayrhofer, but with pale grey to brown,
muriform, 6–9(–10) × (2–)3-septate ascospores, that are (24–)30–34–38(–46) × (11.1–)11.8–13.5–15.2(–
17.8) µm.
TYPE: U.S.A. KANSAS. DOUGLAS CO.: ca. 0.25–0.45 mi N, 1.75 mi E jct of DG Co Rd 1055
& US Hwy 56 in Baldwin City, property of Ralph & Roma Earles, 38.78486°–38.78933°N 95.15201°–
95.15531°W, elev. 970–1060 ft., complex of moderately disturbed, open mixed oak-hickory forest and
woodlands on dissected, N, S and W-facing slopes above tributary to southern arm of Douglas Co. State
Lake; soil sandy, with sandstone outcrops locally common in draws and on middle slopes and limestone
outcrops and boulders occasional on lower slopes, on sandstone along draw on S-facing slope, 18.v.2016,
C.A. Morse 25135 (KANU 399094!, holotype).
DESCRIPTION. – Thallus epilithic, thin (essentially chasmolithic on coarse-grained sandstones) to
rather thick, membranaceous to areolate, sublustrous, greenish grey to pale grey or tan (yellowish in older
specimens), surface smooth to finely verruculose; medulla K/I+ pale blue. Photobiont chlorococcoid, the
cells 7–10(–15) μm in diameter. Ascomata dispersed or in groups of 2–3, borne in hemispherical verrucae;
verrucae (0.2–)0.3–0.4(–0.7) mm in diameter, concolorous with thallus to more commonly contrasting and
dark greyish or yellowish brown with darker brown ostiole, smooth to rough or warted, inspersed with a
few large, colorless to brownish, ± rectilinear crystals (these POL+, persisting in K); involucrellum absent.
Centrum 0.2–0.3 mm in diameter, exciple hyaline below, pale (reddish to greenish) brown or blue-grey
above, K/I+ pale blue; periphysoids simple or sparsely branching, ca. 30 × 1.5 μm; paraphyses thin, ca. 1
µm wide, abundantly branching. Asci narrowly pyriform to cylindrical, ca. 113–164 × 28–36 μm, with (4?)
8 ascospores, KI–, with a distinct ocular chamber. Ascospores ellipsoid to elongate-ellipsoid, or
occasionally broadly clavate, pale grey to brown early in ontogeny (brown when overmature), muriform,
with 6–9(–10) transverse septa and (2–)3–4 ± parallel longitudinal septa, (24–)30–34–38(–46) × (11.1–)11.8–
13.5–15.2(–17.8) µm, L/W ratio = (2.1–)2.2–2.5–2.8(–4.2) [n = 79]. Conidia not observed.
CHEMISTRY. – Spot tests of thallus with K, C, P all negative; thallus UV-. No substances detected
by TLC.
27
Figure 2. Distribution of Thelenella calcicola (A) and T. nubifera (B); inset map: continental U.S.A.
showing the states from which the new species are known. Dashed lines indicate TNC (2007) ecoregions (1
= Central Tallgrass Prairie, 2 = Osage Plains/Flint Hills Prairie, 3 = Cross Timbers, 4 = Ozarks, 5 = South
Atlantic Coastal Plain).
ETYMOLOGY. – The epithet nubifera is Latin (from nubis cloud+ fero to bear) and refers to the
distinctive, grey or brownish spores, which are evocative of little rain clouds.
ECOLOGY AND DISTRIBUTION. – Thelenella nubifera is known from the Central Tallgrass Prairie
and Osage Plains/Flint Hills Prairie ecoregions of eastern Kansas and western Missouri, the northern Cross
Timbers ecoregion in northern Oklahoma, the Missouri Ozarks, and the South Atlantic Coastal Plain
ecoregion in Georgia (TNC 2007; figure 2B). It occurs on shaded or exposed chert and non-calcareous
sandstone, growing on cobbles, boulders, outcrops, and cliffs from 195 to 1050 feet (60–320 meters) above
sea level. Specimens from the eastern Ozarks were collected in mesic, wooded ravines and from shaded
cliffs, while specimens from Kansas, western Missouri, and Oklahoma were found in open oak woodlands
and in sandstone glades. The Georgia collections came from the Altamaha Grit Formation of the Broxton
Rocks area. Whether the species’ presence there may be taken as an indication that T. nubifera is more
broadly distributed throughout the southeastern U.S.A. is unclear, as the Broxton Rocks are known to
harbor disjunct floristic elements from the Appalachian Mountains (Griffin et al. 1994). In Kansas and
Oklahoma, T. nubifera frequently occurs with Fellhanera crucitignorum, and may have similar ecological
requirements through the Cross Timbers (Morse & Ladd 2013). Other associated species include
Anisomeridium distans, Arthonia lapidicola, Bacidina delicata, Endocarpon pallidulum, Fellhanera silicis,
Ionaspis alba, Lepraria finkii, T. brasiliensis, Thelidium minutulum, Trapelia placodioides, Rinodina
oxydata, and Rinodina siouxiana.
DISCUSSION. – Orange et al. (2009) have observed that the medulla of T. muscorum (Fr.) Vainio
var. muscorum is K/I+ pale blue, noting the utility of this character in identification of sterile material. This
reaction was observed in both T. muscorum var. muscorum and T. nubifera, as well as in specimens of T.
brasiliensis, T. modesta, and an unnamed species of Thelenella (Thelenella sp. 1 below), examined for this
study. Mayrhofer (1987) found the presence of amyloid (“J+”) medullary tissue to be unique to T. elliottii
Vainio, Harada (1999) described the thallus of T. luridella to be K/I-, and a K/I+ blue reaction was not
observed in specimens of T. calcicola and T. pertusariella examined for this study. However, the presence
of hemiamyloid tissue may be more common in the genus than published reports suggest.
Pigmented ascospores have been reported from only three other species in Thelenella. Thelenella
hassei (Zahlbr.) H. Mayrhofer, a corticolous species that occurs in coastal southern California, is otherwise
very similar to T. nubifera, differing principally in its ecology and in producing ascospores with only 2–3-
28
longitudinal septa (Mayrhofer 1987). A second corticolous species, T. melanospora Etayo & H. Mayrhofer
documented from the Mediterranean region (Israel, Italy, Spain), produces smaller ascospores (18–22(–27)
× 9–12 μm fide Etayo & Mayrhofer 2003). Thelenella fernandeziana (Zahlbr.) H. Mayrhofer, a saxicolous
species on felsic basalt known from the Juan Fernandez Islands in Chile, produces broader ascospores (15–
22 μm wide fide Mayrhofer 1987).
Although they are invariably pigmented pale grey to brown in the ascus, it is possible to overlook
the pigmentation in very pale ascospores of the new species. In these cases, the pigment is best observed in
optical section of the cell walls, or at lower magnifications (100–400×; see figure 1H). Moreover,
ascospores of Thelenella nubifera often appear colorless very early in ontogeny, and careful examination of
fully mature ascospores may be required to distinguish the species from T. luridella. (Indeed, several
specimens cited here were initially determined as belonging to the latter species.) Ascospores of T. luridella
are also ellipsoid to elongate-ellipsoid and broadly overlap with T. nubifera in size (30–45 × 12–19 μm fide
Mayrhofer 1987), with 7–9 transverse septa and 3 or more longitudinal septa. Thelenella luridella has a
tropical and humid subtropical distribution and has been reported from the Caribbean (Dominica, Trinidad),
South America (Bolivia, Brasil), Arabian Peninsula (Socotra), southern Africa (Lesotho, South Africa),
Asia (Japan, Hong Kong, Nepal), and New Zealand (Mayrhofer 1987, Mayrhofer & McCarthy 1991,
Aptroot & Seaward 1999, Harada 1999, Baniya et al. 2010, McCarthy 2014). The species was reported for
North America by Harris (1995), based on vouchers from Georgia, and subsequently from the Ozarks by
Harris and Ladd (2005). However, these specimens are referred here to T. nubifera. Although one specimen
determined as T. luridella (Mississippi. Wilkinson Co.: Harris 11488 [NY]) was not located, and thus not
available for review, that species is tentatively excluded from North America.
In the field, Thelenella nubifera is most likely to be confused with T. brasiliensis, which appears
to be the more common species in the region, frequently occurs in the same communities and on the same
substrate types. In addition to producing smaller, colorless ascospores, T. brasiliensis may be distinguished
from the new species by its smooth verrucae, which are largely concolorous with the thallus, brownish only
in the immediate vicinity of the ostiole. In T. nubifera, the darkly pigmented area around the ostiole is
typically more extensive. In some specimens of T. nubifera examined (e.g., Morse 24439, see figure 2G),
this pigmented area comprises the greater part of the verrucae, making them contrast strongly with the
surrounding thallus.
Additional specimens examined. – U.S.A. GEORGIA. COFFEE CO.: Broxton Rocks Ecological
Preserve, 9 mi NE of Broxton, 3 mi S of Ocmulgee River, 31°44′N 82°45′W, 16–17.xii.1993, R.C. Harris
32595 (NY), R.C. Harris 32596 (NY). JEFF DAVIS CO.: 0.4 mi E of Coffee County line on Georgia Hwy
107, ca. 11 mi NE of Broxton, ca. 2 mi S of Ocmulgee River, 31°45′N 82°43′W, 5.ii.1995, W.R. Buck
27509 (NY). KANSAS. CHEROKEE CO.: Spring River Wildlife Area, N of NE 100th St, 0.25 mi N of
Old US 96, 37°10′53″N 94°38′58″W, 13.iv.2004, W.R. Buck 46423 (NY), R.C. Harris 48885 (NY).
DOUGLAS CO.: ca. 0.25–0.45 mi N, 1.75 mi E jct of DG Co Rd 1055 & US Hwy 56 in Baldwin City.
property of Ralph & Roma Earles, 38.79°N 95.15°W, 26.ii.2012, C.A. Morse 23280 & K.J. Morse
(KANU), 24.vi.2016, C.A. Morse 25177 (KANU); 1.75 mi N, 0.5 mi W of jct of US Hwy 56 & Co Rd
1055 in Baldwin City, University of Kansas Ecological Reserves: Breidenthal Biological Reserve, 38.81°N
95.20°W, 24.x.2010, C.A. Morse 22125 & K. Logan (KANU), 14.iii.2016, C.A. Morse 25090 (KANU);
1.75 mi N, 0.75 mi W of jct of US Hwy 56 & DG Co Rd 1055 in Baldwin City, private land just N of
University of Kansas Ecological Reserves Wall Woods, 38.81°N 95.20°W, 8.ii.2015, C.A. Morse et al.
24439 (KANU, NY), C.A. Morse et al. 24443 (KANU), C.A. Morse 25117 (KANU), C.A. Morse 25123
(KANU). MONTGOMERY CO.: 0.5 mi N, 5.5 mi W Liberty, S side of Montgomery Co State Lake,
37.16°N 95.70°W, 8.xi.2006, C.A. Morse 14407 (KANU). MISSOURI. DADE CO.: Bona Glade Natural
Area, along MO 215 just E of bridge over Maze Creek Arm of Stockton Lake, 37°32′43″N 93°41′30″W,
16.iv.2005, W.R. Buck 48651 (NY). GENEVIEVE CO.: Magnolia Hollow Conservation Area, along White
Sands Road, 5.5 mi NE of Hwy V, Bloomsdale Quad, 38°02′13″N 90°08′00″W, 30.iii.2006, J.C. Lendemer
et al. 6741 (PH). JEFFERSON CO.: Don Robinson property, N of Sand Cut Road, ca. 1.2 mi E of Hwy
NN, along tributary of La Barque Creek, “Club Moss Hollow,” 38°23′48″N 90°41′31″W, 24.iii.2006, W.R.
Buck 49647 (NY). MONTGOMERY CO.: Graham Cave State Park, jct of roads to Boat Ramp and Indian
Glade Campground, 38°54′17″N 91°34′47″W, 27.x.2001, W.R. Buck 40597 (NY). OKLAHOMA. OSAGE
CO.: 7 mi N, 3 mi E Barnsdall. Woolaroc Wildlife Preserve: along small stream above Swan Lake,
36.66°N 96.11°W, 7.iv.2007, C.A. Morse 14697b & D. Ladd (KANU).
29
Thelenella sp. 1.
Another, evidently undescribed, species is known from the Great Plains, collected once as a small
admixture in northwest South Dakota. In the field, it is most likely to be confused with a species of
Staurothele or Verrucaria, and thus is likely to be overlooked. A provisional description is provided below,
but formal recognition of this taxon awaits additional collections.
DESCRIPTION. – Thallus epilithic, rather thick, areolate, pale grey; medulla inspersed with large
crystals, K/I+ pale blue. Photobiont chlorococcoid, the cells ca. 7.5–14 µm in diameter. Ascomata mostly 1
per areole, immersed in the thallus; ostiole greenish brown, brown in K; involucrellum lacking. Centrum
ca. 0.45 mm in diameter; exciple hyaline (except at ostiole), thin below, thickened to ca. 100 μm thick
above, periphysoids unbranched; paraphysoids ca. 1 μm wide, branching. Asci cylindrical, ca. 73–84 × 21–
23 μm, with (4?)8 ascospores, KI–, without a distinct ocular chamber. Ascospores narrowly ellipsoid to
ellipsoid or broadly fusiform, colorless, muriform, 5–7 × 1–3-septate, ca. 20–33 × 8.5–14 μm. Pycnidia not
observed.
ECOLOGY AND DISTRIBUTION. – Thelenella sp. 1 is known from a single small specimen collected
from non-calcareous siltstone in an open ponderosa pine (Pinus ponderosa) forest on moderate to steep,
west-facing slope with brushy understory at about 3420 feet (1042 meters) above sea level.
DISCUSSION. – Thelenella sp. 1 is unusual in producing a thick, areolate, ashy grey thallus, and
appears to be ecologically distinct from other saxicolous members of the genus. The species should be
compared with T. inductula (Nyl.) H. Mayrhofer, which has been documented from coastal southern
California and eastern Arizona, as well as Macaronesia and Portugal, and which differs in producing an
ochre to light reddish brown thallus (Mayrhofer 1987, 2002b). While other species examined for this study
were found to be at least very pale K/I+ blue, iodine reactivity has been reported elsewhere in the genus
only in the muscicolous species T. muscorum var. muscorum (Orange et al. 2009), and in T. elliottii, which
Mayrhofer (1987) described as having a “J+” medulla. Thelenella elliottii is known only from the
Caribbean (St. Vincent) and differs from the Great Plains specimen in producing a thin, rimose-areolate
thallus and ascospores with 3–4 longitudinal septa (Mayrhofer 1987).
Specimen examined. — U.S.A. SOUTH DAKOTA. HARDING CO.: ca. 3.5 mi S, 18 mi E of
Buffalo, Custer National Forest: East Half: Slim Buttes: Reva Gap: vicinity of Reva Gap Campground,
below The Castles, 45.53°N 103.18°W, 10.vii.2009, C. A. Morse 19635b (KANU).
NEW DISTRIBUTIONAL RECORDS FOR THELENELLA IN THE GREAT PLAINS
In addition to the taxa described above, ongoing fieldwork has yielded a number of noteworthy
distributional records for Thelenella in the southern Great Plains. These are briefly enumerated below.
Thelenella brasiliensis has been reported by Harris (1995) from New Jersey and Oklahoma, and
by Harris and Ladd (2005) from the Ozarks (based on records from Arkansas, Missouri, and northeast
Oklahoma). These are the first reports for Kansas, southwest Oklahoma, and Texas. Specimens were
collected from cobble, boulders, outcrops, and cliffs of non-calcareous rock (fine to coarse sandstones,
granite, and Sioux Quartzite) in sheltered situations in mixed-grass and tallgrass prairie, open oak-hickory
woodlands, and sandstone glades.
Specimens examined. — U.S.A. KANSAS. DOUGLAS CO.: 1.75 mi N, 0.25–0.5 mi W of jct of
US Hwy 56 & DG Co Rd 1055 in Baldwin City, University of Kansas Ecological Reserves: Breidenthal
Biological Reserve, 38.81°N 95.19°W, 17.x.2014, C.A. Morse 24337 & A. Glauser (KANU), 14.iii.2016,
C.A. Morse 25079 (KANU); 4.5 mi S, 2.25 mi W of Stull, Clinton Lake Wildlife Area: above Coblenz
Marsh, 38.90°–38.91°N 95.50°–95.49°W, 28.x.2012, C.A. Morse et al. 23836 (KANU). ELLSWORTH
CO.: 4 mi S, 2 mi E Carneiro, Kanopolis State Park: Horsethief Canyon area, 38.67°–38.68°N 98.00°W,
27.iv.2006, C.A. Morse 12691 & C.C. Freeman (KANU). MONTGOMERY CO.: 0.5 mi N, 5.5 mi W
Liberty. S side of Montgomery Co. State Lake, 37.16°N 95.70°W, 8.xi.2006, C.A. Morse 14392b (KANU).
WOODSON CO.: 6 mi E of Toronto, Woodson County State Fishing Lake & Wildlife Area on SE side,
37.79°N 95.84°W, 28.ix.2008, C.A. Morse 18057b & K. Logan (KANU), 3.ix.2011, C.A. Morse 23100 &
K.J. Morse (KANU). OKLAHOMA. COMANCHE CO.: ca. 4.25 mi N, 2–2.5 mi W of Cache, Wichita
30
Mountains Wildlife Refuge: N side of Eagle Mountain and vicinity of The Narrows, 34.70°N 98.67°–
98.68°W, 14–15.iv.2010, C.A. Morse 20515c (KANU). OKMULGEE CO.: 0.5–0.75 mi N, 6 mi W of jct of
US Hwys 62 & 75 on S side of Okmulgee, Okmulgee State Park, on peninsula just N of Blackjack Area
tent camping sites, 35.62°N 96.07°W, 6.ix.2009, C.A. Morse 20203c & K. Logan (KANU). TEXAS.
PARKER CO.: ca. 2 mi N, 4.5 mi E jct of US Hwys 180W and 281 in Mineral Wells, Lake Mineral Wells
State Park: NW side of lake, just N and E of Cross Timbers Camping Area, 32.83°N 98.04°W, 26.iv.2009,
C.A. Morse 18491b & D. Ladd (KANU).
Thelenella modesta was reported by Harris (1995) from California and Minnesota and by Tucker
and Egan (2009) from Texas. Ascospores examined for this study were (19–)22–29(–37) × (8–)8.5–13(–16)
μm, slightly smaller than previously reported (25–42 × 11–17 μm fide Mayrhofer 1987). These are the first
reports for Kansas, where the species was collected from distal branches of green ash (Fraxinus
pennsylvanica) and redbud (Cercis canadensis) in upland tallgrass prairie remnants, and for North Dakota,
where the species was found on aspen (Populus tremuloides) and a large cottonwood (P. deltoides).
Specimens examined. — U.S.A. KANSAS. BOURBON CO.: ca. 4 mi S, 5.5 mi W of Uniontown,
SW side of Bourbon Co. State Fishing Lake and Wildlife Area, 37.79°N 95.08°W, 11.xii.2015, C.A. Morse
24948 (KANU); 3 mi S, 0.5 mi E of Fulton, Unique Prairie, 37.96°N 94.70°–94.71°W, 2.iv.2016, C.A.
Morse et al. 25098 (KANU). NORTH DAKOTA. ROLETTE CO.: 5.5 mi W of St. John, Wakopa
Wildlife Management Area, 48.95N 99.83W, 24.viii.2015, M.K. Advaita 17897-B (KANU). TOWNER
CO.: 0.5 mi E, 1.5 mi N of Cando, U.S. Waterfowl Production Area, 48.51N 99.19W, 22.viii.2015, M.K.
Advaita 17839 (KANU).
Thelenella muscorum var. muscorum was reported by Harris (1995) and Fryday and Coppins
(2004) from Colorado, Minnesota, and North Carolina, by Mayrhofer (2002a) from Arizona, California,
and northwestern Mexico, and the Ozarks (Harris & Ladd 2005, D. Ladd personal comm.). The eight-
spored variety, T. muscorum var. octospora (Nyl.) Coppins & Fryday appears to be restricted to the western
North America, with reports from Colorado, Idaho, Montana, Oregon, and Saskatchewan (Anderson 1962,
McCune et al. 2014). Thelenella muscorum var. muscorum is new for Kansas and Oklahoma. All
specimens were collected on bryophytes overgrowing sandstone in open oak-hickory woodlands.
Specimens examined. — U.S.A. KANSAS. DOUGLAS CO.: ca. 0.25–0.45 mi N, 1.75 mi E jct of
DG Co Rd 1055 & US Hwy 56 in Baldwin City, property of Ralph & Roma Earles, 38.78°N 95.15°W,
18.v.2016, C.A. Morse 25147 (KANU); 1.75 mi N, 0.5 mi W of jct of US Hwy 56 & Co Rd 1055 in
Baldwin City. University of Kansas Ecological Reserves: Breidenthal Biological Reserve, 38.81°N
95.19°W, 24.x.2010, C.A. Morse 22122 & K. Logan (KANU), 14.iv.2015, C.A. Morse 24470a & A.
Glauser (KANU). CHAUTAUQUA CO.: 3 mi N, 1 mi W of Hale, West Liberty Cemetery and environs,
37.28°N 96.04°W, 7.ix.2009, C.A. Morse 20256b & K. Logan (KANU). FRANKLIN CO.: 3 mi S, 1 mi E
Homewood., Ottawa University Natural History Reservation (“Ferndale”), 38.47°N 95.36°W, 5.iii.2008,
C.A. Morse 16341a & K. Logan (KANU). LINN CO.: 2.75 mi W of jct of KS Hwys 7 & 52 in Mound City,
Dingus Natural Area, 38.13°N 94.87°W, 31.i.2009, C.A. Morse 18236 & K. Logan (KANU). WOODSON
CO.: 6 mi E of Toronto, Woodson Co. State Fishing Lake & Wildlife Area on SE side, 37.79°N 95.84°W,
28.ix.2008, C.A. Morse 18036 & K. Logan (KANU), 18.iii.2012, C.A. Morse 23327 & K.J. Morse
(KANU). OKLAHOMA. OSAGE CO.: 7.5 mi N, 3 mi E Barnsdall, Woolaroc Wildlife Preserve: start of
nature trails area, just NW of museum complex, 36.67°N 96.11°W, 6.iv.2007, C.A. Morse 14632b & D.
Ladd (KANU); 7.5 mi N, 3.75 mi E Barnsdall, Woolaroc Wildlife Preserve: just W of Little Rock Creek, N
of outlet of Clyde Lake, 36.67°N 96.10°W, 7.iv.2007, C.A. Morse 14802a & D. Ladd (KANU).
Thelenella pertusariella was reported by Harris (1995) from Michigan and Minnesota, and by
Harris and Ladd (2005) from Oklahoma. Harris (1995) noted that North American specimens produced
ascospores with more septa than were found in European material. Ascospores examined for this study did
not differ significantly from the description by Mayrhofer (1987), with 5–10 transverse septa and
infrequent longitudinal septa (with only one or two cells in each ascospore divided longitudinally).
However, ascospores were (16–)19–26(–32) × 5.5–7.5 μm, slightly narrower than reported for the species
by Mayrhofer (1987) (22–35 × 7–10 μm) or Harris (1995) (23–34 x 7–11 μm). These are the first reports
for Kansas. Both specimens were collected from oaks (Quercus) in mesic, mixed oak-hickory(-basswood)
forest.
31
Specimens examined. — U.S.A. KANSAS. DOUGLAS CO.: 1.75 mi N, 0.5 mi W of jct of US
Hwy 56 & DG Co Rd 1055 in Baldwin City, University of Kansas Ecological Reserves: Breidenthal
Biological Reserve, 38.81°N 95.19°W, 8.ii.2009, C.A. Morse 18256 (KANU). LEAVENWORTH CO.:
Fort Leavenworth Military Reservation: W-central part, area roughly bounded by ravine 0.25 mi E of Bell
Point, between reservoir (to N) and radio tower (to S), and area E of reservoir and below (E of) Hancock
Hill, in vicinity of Girl and Boy Scout Camps, 39.36°–39.37°N 94.93°W, 14.v.2008, C.A. Morse 16480b
(KANU).
WORLDWIDE KEY TO THELENELLA
When measuring ascospores, users should take care to base their observations on mature
ascospores, as immature ascospores are frequently smaller than the values reported in published accounts,
and overmature ascospores are frequently much narrower. Further, because ascospore size is critical in
determining many species of Thelenella, it is important (if often difficult!) to measure a sufficient number
of ascospores to ascertain the typical size range. In order to avoid confusion regarding septation and size,
counts of the number of transverse and longitudinal septa are separated by a slash [ / ], while indications of
length and width are separated here by a multiplication sign [ × ]. Thus, a species with muriform ascospores
with 7 to 10 transverse septa and 3 to 4 longitudinal septa will be indicated as “ascospores 7–10 / 3–4-
septate”. Species with transversely septate to submuriform ascospores are indicated in the same way, so it
is important to bear in mind that “ascospores 7–10 / 0–2–septate” means that some ascospores may be
transversely septate only. Measurements and distributional information were taken principally from
Mayrhofer (1987) and Harris (1995), and supplemented by descriptions in Santesson (1952), Mayrhofer
and Poelt (1985), Mayrhofer and McCarthy (1991), Kalb (1995), Harada (1999), Seaward and Aptroot
(2000), Etayo and Mayrhofer (2003), Fryday and Coppins (2004), Pinokiyo and Singh (2006), Lücking
(2008), Kinalioglu and Aptroot (2011), Aptroot et al. (2014), and McCune et al. (2014). Distributions are
specified to the best of the author’s ability, especially for seldom-collected taxa and for species with
documented ranges in North America. Unfortunately, many taxa remain very poorly known, so both
measurements and distributional information must be regarded with caution. Some distinctions between
species remain uncertain, and additional fieldwork is needed to characterize the range of morphological and
ecological variability among members of the genus.
1. Muscicolous ............................................................................................................................................... 2
2. Ascospores 7–8 / 0–1-septate, 22–30 × 7–10 µm; Europe (Austria) .....................................................
............................................................................... T. vezdae (H. Mayrhofer & Poelt) Coppins & Friday
2. Ascospores richly muriform, > 40 × 15 µm ....................................................................................... 3
3. Ascospores 2–4 per ascus, ca. 21 / 2–7-septate, 60–100 × 20–27 µm; northern Africa (Tunisia),
Europe (broadly distributed), North America (U.S.A.: broadly distributed) .......................................
.......................................................................................... T. muscorum (Fr.) Vainio var. muscorum
3. Ascospores 6–8 per ascus, 11–14 / 1–3-septate, 40–60 × 15–20 µm; northern Europe, North
America (Canada: Saskatchewan, northwest U.S.A.: Idaho, Oregon) ...............................................
...................................................................... T. muscorum var. octospora (Nyl.) Coppins & Fryday
1. Not muscicolous (corticolous, foliicolous, or saxicolous) ......................................................................... 4
4. Saxicolous .......................................................................................................................................... 5
5. Ascospores > 30 µm long (species with largest ascospore 32 [36] µm long key both ways) ..... 6
6. Involucrellum present ........................................................................................................... 7
7. Ascospores 5–7 / 3-septate, 20–32 × 10–16 µm; Subantarctic islands (Kergulen,
Marion, Heard islands)................................................. T. kerguelena (Nyl.) H. Mayrhofer
7. Ascospores > 7 / 3-septate, > 34 × 14 µm ..................................................................... 8
8. Thallus pale greenish grey; ascospores 12–16 / 3–4 septate, 34–52 × 14–20 µm;
Antarctic islands (Bouvet and South Georgia islands), Subantarctic islands
(Kerguelen, Heard, Macquaire islands) .........................................................................
..................................... T. mawsonii (C.W. Dodge) H. Mayrhofer & P.M. McCarthy
8. Thallus ochre to light grey-brown; ascospores 7–11 / 3 septate, 35–47 × 14–19
µm; western North America (Mexico: Baja California, U.S.A.: California) .................
................................................................................................ T. weberi H. Mayrhofer
6. Involucrellum absent ............................................................................................................ 9
32
9. Ascospores pigmented grey or brown early in ontogeny ............................................ 10
10. Thallus light brown to grey brown; ascospores 7–9 / 2–5?-septate, 30–45 × 15–
22 µm; on felsic basalt; western South America (San Juan Fernandez Islands) ...........
..................................................................... T. fernandeziana (Zahlbr.) H. Mayrhofer
10. Thallus tan to greenish grey; ascospores 6–9(–10) / 3–4-septate, (24–)30–38(–46)
× (11.1–)11.8–15.2(–17.8) µm; on sandstone; southeast North America (U.S.A.:
Kansas, Missouri, Oklahoma, Georgia) ................................. T. nubifera C.A. Morse
9. Ascospores colorless (although overmature ascospores may become brown) ............ 11
11. Ascospores transversely septate to submuriform ............................................... 12
12. Ascospores 11–14 / 0–2 septate, 44–60 × 10–14 µm; on siliceous rocks;
western Europe (United Kingdom) ........................................................................
...................................................... T. larbalestieri (A.L. Sm.) Coppins & Fryday
12. Ascospores (5–)7–11(–12) / 1(–2)-septate, (18.5–)21–29(–33) × (5.4–)6.3–
8.4(–9.8) μm; on calcareous rocks; central North America (U.S.A.: Kansas,
Missouri)......................................................................... T. calcicola C.A. Morse
11. Ascospores muriform, with 3 or more longitudinal septa .................................. 13
13. Ascospores broadly ellipsoid, 7–11 / 3–4–septate, 20.5–35.5 × 12–18.5 μm;
Oceania (southern Australia: Bass Strait Islands, Tasmania) ................................
..................................................... T. tasmanica H. Mayrhofer & P.M. McCarthy
13. Ascospores elongate-ellipsoid to subcylindric to spindleform, > 30 μm long .
........................................................................................................................... 14
14. Ascospores with 7–9 / 3+-septate, 30–45 × 12–19 μm; southern Africa
(Lesotho, South Africa), southern Asia (Arabian Peninsula: Socotra, Japan,
Hong Kong, Nepal), Caribbean (Dominica, Trinidad), southern South
America (Bolivia, Brazil), southeast Oceania (New Zealand) ........................
......................................................................... T. luridella (Nyl.) Mayrhofer
14. Ascospores with 9–17 / 3–5-septate, > 40 µm long ............................. 15
15. Ascocarp 0.2–0.4 mm in diameter; ascospores 9–13 / 3-septate, 40–
55 × 14–20 µm; southern Asia (Indonesia: Java) ....................................
..................................................... T. marginata (Groenh.) H. Mayrhofer
15. Ascocarp 0.5–1 mm in diameter; ascospores 11–17 / 3–5-septate,
50–82 × 22–33 µm; Antarctic (South Shetland Islands) .........................
......................................................... T. antarctica (M. Lamb) Ericksson
5. Ascospores < 30 (36) µm long .................................................................................................. 16
16. Involucrellum present; ascospores 5–7 / 3-septate, 20–32 × 10–16 µm; Subantarctic
islands (Kergulen, Marion, Heard islands) ........................................................... T. kerguelena
16. Involucrellum absent ........................................................................................................ 17
17. Ascospores transversely 3-septate, 18–23 × 6.5–7.5 µm; Caribbean (Puerto Rico) .......
.................................................................... Thelenella sp. Buck 18291A of Harris (1995)
17. Ascospores submuriform or muriform ...................................................................... 18
18. Ascospores irregularly submuriform, (5–)7–11(–12) / 1(–2)-septate, (18.5–)21–
29(–33) × (5.4–)6.3–8.4(–9.8) μm; on calcareous rocks; central North America
(U.S.A.: Kansas, Missouri) ....................................................................... T. calcicola
18. Ascospores muriform, with 2 or more longitudinal septa; on non-calcareous
rocks ......................................................................................................................... 19
19. Ascospores 6–7 / 2–3 septate, 18–20 × 8–11 µm; eastern North America
(U.S.A.: New York) .......................................................... T. humilis R. C. Harris
19. Ascospores 20–30 (36) µm long ................................................................. 20
20. Ascospores broadly ellipsoid, 20.5–35 × 10–18.5 μm wide, with 3–4
longitudinal septa ...................................................................................... 21
21. Thallus pale to dark greenish grey to grey-brown; ascospores 7–11 /
3–4–septate, 20.6–35.3 × 11.8–18.5 μm; Oceania (southern Australia:
Bass Strait Islands, Tasmania) ........................................... T. tasmanica
21. Thallus whitish to yellowish grey; ascospores 6–7 / 3–4 septate,
22–27 × 10–14 µm; on volcanic rock; Caribbean (St. Vincent) ..............
..................................................................................... T. elliottii Vainio
33
20. Ascospores elongate-ellipsoid, < 8–13 µm wide, with 1–3 longitudinal
septa ........................................................................................................... 22
22. Ascospores 6–9 / 1–2 septate, 20–27 × 8–11 μm, with ± pointed
ends; Caribbean (Puerto Rico), southern North America (U.S.A.:
Louisiana) ......................................................... T. sastreana R.C. Harris
22. Ascospores 5–8 / 1–3 septate, 9–14 μm wide, with ± rounded ends .
............................................................................................................. 23
23. Thallus thick, rimose-areolate to areolate ............................. 24
24. Thallus ochre to pale reddish brown; ascospores 6–8 /
(1)2–3+-septate, 24–36 × 9–13 μm; on lava, granite, schist,
slate; southwest Europe (Portugal), Macaronesia, southwest
North America (U.S.A.: Arizona, California) ...........................
............................................. T. inductula (Nyl.) H. Mayrhofer
24. Thallus ashy grey; ascospores 5–7 / 1–3-septate, 20–32 ×
9–14 µm; on non-calcareous siltstone; central North America
(U.S.A: South Dakota) ............... Thelenella sp. 1 of this paper
23. Thallus thin, membranaceous to rimose ................................ 25
25. Thallus pale ochre to yellow-brown, membranous-rimose,
matt; ascomata 0.5–0.8 mm in diameter; ascospores 5–7 / 2–3-
septate, 24–30 × 9–13 μm; northern Europe (Svalbard),
northeast North America (Canada: Ellesmere Island;
Greenland: Disko Island) ...........................................................
......................................... T. sordidula (Th. Fr.) H. Mayrhofer
25. Thallus pale brown, olive-brown, or olive-green,
membranous to membranous-rimose, matt to glossy; ascomata
0.2–0.4 mm in diameter; ascospores 6–7 / 2+-septate, 20–32 ×
9–13 μm; western Africa, Asia (China?), Central America
(Costa Rica), South America (Brazil), eastern North America
(U.S.A.: Arkansas, Kansas, Missouri, New Jersey, Oklahoma,
Texas) .................................. T. brasiliensis (Müll. Arg.) Vainio
4. Species corticolous or foliicolous ..................................................................................................... 26
26. Species corticolous .................................................................................................................. 27
27. Involucrellum present; ascospores transversely 11–19-septate, fusiform, 50–100 ×
14–19 µm; Central America (Costa Rica, Panama), southern North America (Mexico:
Veracruz, U.S.A.: Florida), South America (Brazil, Paraguay) ...........................................
... T. geminipara (Malme) R. C. Harris [= Aspidothelium geminiparum (Malme) R. Sant.]
27. Involucrellum absent (although ascoma wall reported to be carbonized in A.
lateralis); ascospores irregularly transversely septate, submuriform to muriform
(transversely septate to submuriform in A. lateralis), < 60 µm long .............................. 28
28. Ascospores becoming grey or brown early in ontogeny, muriform with 2–4
longitudinal septa ..................................................................................................... 29
29. Ascospores 25–40 × 10–17 µm; southwest North America (U.S.A.:
California) ......................................................... T. hassei (Zahlbr.) H. Mayrhofer
29. Ascospores 18–22(–27) × 9–12 µm; Mediterranean (Israel, Italy, Spain) ........
................................................................ T. melanospora Etayo & H. Mayrhofer
28. Ascospores colorless (although overmature ascospores may become brown),
septation various ...................................................................................................... 30
30. Ascospores with 11–15 transverse septa, 35–60 μm long .......................... 31
31. Ascospores submuriform, with 1–2 longitudinal septa, 35–50 × 9–12
μm, with pointed ends; southern Europe (Greece) .........................................
.......................................................... T. justii (Servít) H. Mayrhofer & Poelt
31. Ascospores muriform, with 3 longitudinal septa, 42–60 × 13–19 μm,
with rounded ends; southern Asia (Indonesia), South America (Paraguay) ...
................................................................................ T. paraguayensis Malme
30. Ascospores with 6–10 transverse septa, < 42 μm long ............................... 32
34
32. Ascospores muriform, with 1–4 longitudinal septa, and > 9 μm wide ......
................................................................................................................... 33
33. Thallus greyish green; ascospores ellipsoid, 6–8 / 1–3-septate, 20–
28 × 9–15 μm; Caribbean (Jamaica) ........................ T. follmannii Kalb
33. Ascospores 7–9 / 2–4-septate, > 25 × 11 μm ................................ 34
34. Thallus yellowish grey, greyish brown, or brownish; asci
cylindrical to clavate; ascospores ellipsoid to elongate-ellipsoid,
with 2–3 longitudinal septa, 25–42 × 11–17 μm; northern Africa
(Morocco), Asia (Israel, Turkey), Australia, Europe (broadly
distributed), Indian Ocean (Chagos Archipelago), western North
America (U.S.A.: California, Kansas, Minnesota, North Dakota,
Texas) .......................................................... T. modesta (Nyl.) Nyl.
34. Thallus whitish to olivaceous; asci obclavate to pyriform;
ascospores ovoid, with 2–4 longitudinal septa, 30–38 × 13–15 μm;
southeast North America (U.S.A.: Florida) ... T. rappii R. C. Harris
32. Ascospores transversely septate to (irregularly) submuriform, with 0–2
longitudinal septa, and < 11 μm wide ........................................................ 35
35. Ascomata 0.4–0.55 mm in diameter; ostiole lateral; ascospores
7–9 / 0–2-septate, 27–32 × 9–10.5 μm; South America (Brasil) .......
.................................................... T. lateralis Aptroot & M. Cáceres
35. Ascomata < 0.4 mm in diameter; ostiole apical; ascospores
various; distribution various (but not known from South America) ..
...................................................................................................... 36
36. Species corticolous on dead spruce or muscicolous on
dead mosses; thallus whitish or brownish; ascospores 7–8-
transversely septate, with 0–1 longitudinal septa, 22–30 × 7–
10 µm; central Europe (Austria) ............................... T. vezdae
36. Species corticolous; ascospores submuriform to muriform,
with 1–2 longitudinal septa; distribution various ................. 37
37. Thallus indistinct, fleck-like to membranous-cracked,
dirty whitish to yellowish; ascomata 0.2–0.4 mm in
diameter; ascospores 6–10 / 1–2-septate, 22–35 × 7–10
μm [central North American specimens (16–)19–26 (32)
× 5.5–7.5 μm]; Europe (Austria, Finland, Norway, Russia,
Sweden, Switzerland), central North America (U.S.A.:
Kansas, Michigan, Minnesota, Oklahoma) ........................
............................................ T. pertusariella (Nyl.) Vainio
37. Thallus discontinuous to membranous or
membranous-cracked, light brown to light grey; ascomata
0.1–0.3 mm in diameter; ascospores 6–7 / 1–2–septate,
20–30 × 7–11 μm; southwest North America (U.S.A.:
California), Oceania (southeast Australia: New South
Wales) .............. T. sychnogonioides (Zahlbr.) R. C. Harris
26. Species foliicolous .......................................................................................................................... 38
38. Ascospores transversely septate with > 11 septa and > 50 μm long ........................................ 39
39. Involucrellum present; ascospores transversely 11–19-septate, fusiform, 50–100 × 14–19
µm; Central America (Costa Rica, Panama), southern North America (Mexico: Veracruz,
U.S.A.: Florida), South America (Brazil, Paraguay) ........................................... T. geminipara
39. Ascospores 15–29-transversely septate, 50–90 × 12–20 μm; Central America (Costa Rica,
Guatemala, Honduras, Panama), southern North America (Mexico: Oaxaca, Veracruz, U.S.A.:
Florida), South America (Brazil, Ecuador, French Guiana, Guyana, Peru) ................................
............... T. fugiens (Müll. Arg.) R. C. Harris [= Aspidothelium fugiens (Müll. Arg.) R. Sant.]
38. Ascospores submuriform or muriform, with at least 1 longitudinal septum OR (if transversely
septate) < 40 μm long .................................................................................................................... 40
35
40. Ascomata 0.15–0.18 mm in diameter; ascospores 3–7 / 0–2-septate, (16) 25–40 × 7–12
μm; southern Asia (India) ..................................................... T. indica Pinokiyo & Kr.P. Singh
40. Ascomata > 0.2 mm in diameter; ascospores > 8 / 1-septate, > 34 × 12 μm; distribution
various (but not known from India) ....................................................................................... 41
41. Involucrellum present ............................................................................................... 42
42. Ascomata 0.3–0.5 mm in diameter; ascospores 20 × 1–3-septate, 35–70 × 15–25
μm; Central America (Costa Rica, El Salvador), southern North America (U.S.A.:
Florida), South America (Brazil, Ecuador, Peru) ..........................................................
................ T. cinerascens (Vainio) R. C. Harris [=Aspidothelium cinerascens Vainio]
42. Ascomata 0.2–0.3 mm in diameter; ascospores 17–20 × 1–3-septate, 35–45 × 12–
20 μm; Central America (Costa Rica, Guatemala), South America (Ecuador) .............
......................................................... T. trichothelioides (Serús. & Vězda) R.C. Harris
[= Aspidothelium trichothelioides Serús. & Vězda]
41. Involucrellum absent ................................................................................................. 43
43. Ascomata 0.4–0.6 mm in diameter; ascospores 8–17 / 1–4-septate, 35–64 × 13–
20 μm, ellipsoid; northern Oceania (New Guinea) .......................................................
....................................... T. verruculosa (R. Sant.) ined. (see Farkas & Sipman 1997)
[= Aspidothelium verruculosum R. Sant.]
43. Ascomata 0.3–0.4 mm in diameter; ascospores 11–15 / 2–3-septate, 45–65 × 17–
23 μm, elongate to spindleform; western Africa (Tanzania) .........................................
............................................................................. T. fusispora Vězda & H. Mayrhofer
ACKNOWLEDGEMENTS
I thank Doug Ladd (The Nature Conservancy, Saint Louis, Missouri), James Lendemer (NY) and two
anonymous reviewers for their careful reading of the manuscript; Dick Harris (NY) and Helmut Mayrhofer (GRAZ) for
conferring about the identities of the new species; the staff at NY and PH for kindly providing loans of specimens for
study; M.K. Advaita for putting recent collections of T. modesta at my disposal; property owners and managers for
permission to collect; and Deborah Borek, Craig Freeman, Abby Glauser, Courtney Jarvis, Doug Ladd, Katherine and
Helen Morse, and Gary Tegtmeier for assistance and companionship in the field. Fieldwork was made possible by an
anonymous contribution to the McGregor Herbarium Endowment Fund.
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