Type. ARGENTINA. 1889, Lorentz s.n. (G ‑ holotype!).

Rinodina metabolica f. leioplaca Müll. Arg., Rev. Mycol. 10: 63 (1888).
Type. PARAGUAY. Ascuncion, 1878, Balansa s.n. (G ‑ holotype!).

Rinodina haplosporoides Müll. Arg., Bull. Soc. Roy. Bot. Belgique 30: 62 (1891).
Type. COSTA RICA. Baie de Salinas, 1890, H. Pittier 5223 (G ‑ holotype!).

Rinodina prasina Müll. Arg., Bull. Soc. Roy. Bot. Belgique 30: 61 (1891).
Type. COSTA RICA. San José, 1890, H. Pittier 5260 (G ‑ holotype!).

Rinodina applanata H. Magn., Bot. Not. 1947: 43.
Type. [U.S.A.] LOUISIANA. near St. Martinsville, 2nd August, 1895, A.B. Langlois s.n. (U ‑ holotype!, COL ‑ isotype!).

Rinodina neglecta Aptroot, Proc. Kon. Ned. Akad. Wetensch., C90: 240 (1987).
Type. FRENCH GUYANA. Cayenne, Place des Palmistes, on Maximiliana (Palmae), March 1985, Aptroot 15081 (U - holotype!).

Exsiccata. Kalb Lich. Neotrop. 118 (GZU as R. intrusa); Malme Lich. Austroamer. 58A, B (H as R. intrusa); Vězda Lich. Sel. Exs. 1981 (CANL as R. applanata).

Description. Thallus thin, light grey to dark greenish-grey, sometimes with an ochraceous tinge, discontinuous or continuous becoming rimose, or rimose‑areolate; areoles to 0.40-0.50 mm wide, margins sometimes regenerating; surface plane or minutely rugose, the whole thallus then with a somewhat scabrid appearance, matt or shining; margin determinate or not; prothallus absent or thin, dark, entire; vegetative propagules absent.  Apothecia erumpent becoming broadly attached, rarely narrowly attached, frequent, scattered, sometimes contiguous, to (0.30‑)0.45‑1.00 mm in diam.; disc dark brown to black, plane becoming slightly, or rarely, strongly convex; thalline margin sometimes poorly developed, then growing up from the base of apothecium, concolourous with thallus, becoming entire or often crenulate when incomplete, 0.05‑0.10 mm wide, persistent; excipular ring usually present, raised.  Apothecial Anatomy. Thalline exciple 30‑65(‑105) µm wide laterally, sometimes absent; cortex 5‑15 µm wide; epinecral layer sometimes present, 5‑15 µm wide; crystals absent from cortex and medulla; cortical cells to 4.0‑6.0 µm wide, not or hardly pigmented; algal cells to 8.0‑13.0 µm long; thalline exciple (20‑)25‑40(-100) µm wide below; proper exciple usually light brown, often well developed, (5‑)10‑30 µm wide laterally, (10‑)15‑65 µm above; hypothecium hyaline, (30‑)50‑100(‑120) µm deep, tapering into a stipe; hymenium 60‑120 µm high, not inspersed; paraphyses 1.5‑2.5 µm wide, conglutinate or not, apices to 3.5‑5.5 µm wide, lightly pigmented immersed in dispersed pigment forming a light brown to red‑brown epihymenium; asci (45‑75) x (14‑30) µm. Ascospores 8/ascus, Type B development, Pachysporaria‑type II, (12.0-) 17.0‑18.5(-23.5) x (6.5-)8.0‑8.5(-10.0) µm, average l/b ratio 2.0-2.3, often curved when immature, more broadly‑ellipsoid at maturity, lumina often with an irregular or lacrimiform shape during development, becoming ovoid to rounded; torus narrow at maturity; walls lightly pigmented, not ornamented. Pycnidia reddish-brown, often half exposed; conidiophores Type I; conidia bacilliform, 3.0‑4.5 x 1.0-1.5 µm.

Chemistry. Spot tests all negative; secondary metabolites not detected.

Substrate and Ecology. Corticolous, the species has been collected on Acer drummondii, A. rubrum,  Azalea, Baccharis halimifolia, Carpinus caroliniana, Carya illinoensis, C. tomentosa, Celtis laevigata, C. occidentalis, Diospyros, Fagus, Gleditsia triacanthos, Ilex opaca, Juniperus virginiania, Lagerstroemia indica, Leucaena, Lyonia, Lysiloma latisiliqua, Platinus occidentalis, Populus balsamifera, Prunus maritima, P. serotina, Quercus alba, Q. coccinea, Q. ilicifolia, Q. montana, Q. stellata, Q. velutina, Robinia pseudoacacia, Sabal palmetto, Salix, Taxodium, Ulmus crassifolia and Vaccinium, to a maximum recorded elevation of 750 m. Rinodina maculans has been collected once with R. dolichospora.

Distribution. The species has a Coastal Plain distribution in North America being found from Texas to as far north as Massachusetts, also extending up the Mississippi valley to southern Missouri and Illinois. The record for R. applanata reported from Long Island (Brodo 1968) belongs to R. subminuta although R. maculans is present there (reported by Brodo as R. pachysperma). Rinodina maculans is widely distributed in Central and South America (Malme 1902 as R. intrusa).

Notes. Rinodina maculans is characterized by its narrowly ellipsoid, Pachysporaria-type II spores and Coastal Plain distribution. The pachyspermous spores are variable in shape and length (CV >10%). They are at first narrowly ellipsoid, hardly pigmented, and sometimes curved even after release from the ascus. At maturity they become regularly ellipsoid in shape, sometimes broadly so, and remain lightly pigmented. The species has erumpent and persistently broadly attached apothecia and may be closely related to R. intrusa through its prasina-morph discussed below. Rinodina intrusa is distinct in its smaller apothecia, more continuous thallus and particularly in its smaller, broadly‑ellipsoid ascospores. Rinodina colobinoides, another Coastal Plain species from the Gulf states, is distinguished primarily by it blastidiate thallus.

It is unfortunate when a species is based on a type specimen that has an atypical morphology, the more so when its thallus has been compared to bird excrement (Malme 1902)! The very thin, dark brownish thallus and apothecia with predominantly convex discs and lacking thalline margins are an unusual combination of characters. However, the erumpent, young apothecia with poorly developed thalline margins are characteristic of the R. prasina morphotype, as is the more or less continuous thallus with its shining surface. This morph is uncommon in North America where it is apparently restricted to mesic environments in Florida. It is much more common in South America where it can be seen to intergrade with the R. applanata morphotype.

The applanata‑morph is often characterized by a well developed proper exciple and often partly developed or crenulate thalline margin. This is best seen in young apothecia where the thalline margin grows up and around the proper exciple in lobes, frequently resulting in a crenulate margin in older apothecia. The proper exciple often remains as a prominent excipular ring within the fully formed thalline margin. This type of margin is not uncommon in corticolous species but is perhaps most frequently found in species with southern affinities such as R. colobinoides and R. granuligera.

Rinodina maculans is particularly frequent and well developed in the Cape Cod region of Massachusetts and Suffolk Co, New York. It was recorded from the latter by Brodo (1968) as R. pachysperma, who noted the prominent proper margin in macroscopic view. The latter species is distinguished from R. maculans by its more broadly ellipsoid, Pachysporaria-type spores and its Great Lakes-northeastern distribution. Harris (personal communication) indicates that the species is very common in the Ozarks.

Specimens examined. U.S.A. ALABAMA. Marion Co., Hackleburg, E. Lay 99‑938 (personal herb.); Tuscaloosa Co., 15 mi E Tuscaloosa, B.D. Ryan 29272 (ASU); ARKANSAS. Garland Co., Hot Springs Nat. Park, C.M. Wetmore 85899, 86054, 86217, 86319, 86535 (MIN); Music Mountain, C.M. Wetmore 86398 (MIN). FLORIDA. Duval Co., Big Talbot Island, R.C. Harris 23049c, 23918 (NY); Lee Co., Caloosa River, 1878, R.A. Austin (FH); Leon Co., Apalachicola Nat. Forest, R.C. Harris 23268 (NY); Marion Co., 3.5 mi E Cedar Creek, E. Lay 95‑1598 (personal herb.); Miami‑Dade Co., Miami, S.C. Tucker 25286 (LSU); S tip Key Biscayne, S.C. Tucker 25297 (SBBG); Polk Co., Bok Tower Gardens, 1989, E.M. Wheeler (NY); E Lake Wales, C.M. Wetmore 13978 (MIN); Nalcrest, 1988, 1989, E.M. Wheeler (NY); Seminole Co., Sanford, 1911, S. Rapp (FH); Volusia Co., South Daytona, I.M. Brodo 16435 (CANL). GEORGIA. Haines Falls, H.W. Ravenel 36 (FH); Columbia Co., Appling, E. Lay 99‑397 (personal herb.); Glynn Co., Jekyll Island, I.M. Brodo 16441, 16449 (CANL). ILLINOIS. Pope, Co. Lusk Creek, A.C. Skorepa 6224 (BALT). LOUISIANA, Silan, 1895, A.B. Langlois (US); Cameron Par., Peveto Beach, S.C. Tucker 6787 (WIS); E Baton Rouge Par., Baton Rouge, S.C. Tucker 12076 (LSU, MIN); 17407, 25339 (NY); 15675, 25589A (LSU); 7305, 7408, 21135, 26595 (MIN); 11052 (MSC); 15945 (COLO, WIS); 20974b (CANL); Iberville Par., Plaquemine, A.B. Langlois 112 (US); St. Gabriel Church, A.W. Lievens 5394 (SBBG); Plaquemines Par., Pointe a la Hache, 1884, A.B. Langlois (US); Pointe Coupee Par., New Roads, M. Lieux 9995 (MSC); St. Martin Par., Breaux Bridge, A.B. Langlois 855; 1893, A.B. Langlois; St. Martinsville, A.B. Langlois 482 (all US); 1894, A.B. Langlois (NY, US, US); 1895, A.B. Langlois  (COLO, UPS, US); W Felicina Par., Jackson, S.C. Tucker 28210 (MIN, NY); near Thomson Creek, S.C. Tucker 31527 (SBBG). MARYLAND. Wicomico Co., 4 mi SW Delmar, 1975, C. Phillips (LSU); Worcester Co., Milburn Landing State, A.C. Skorepa 11038 (SASK).  MASSACHUSETTS. Barnstable Co., Eastham, Cape Cod, I.M. Brodo 4251 (MSC); Morris Island, I.M. Brodo 4357 (COLO, MSC); 4393 (CANL); near Sandwich, I.M. Brodo 4527 (MSC); Wellfleet, 2000, E. Kneiper (personal herb.); Bristol Co., New Bedford, H. Willey (FH, MIN, US); H. Willey 114 (FH); Dukes Co., Marthas Vineyard, 1994, E. Lay (personal herb.); E. Lay 94‑1577 (personal herb.); Essex Co., West Newbury, 1934, S.K. Harris (MIN); Nantucket Co., 2 mi E Surfside, I.M. Brodo 4092 (MSC); Saul's Hill, I.M. Brodo 3993a (MSC, WIS); Polpis Road, I.M. Brodo 3978 (MSC); Tom Never's Head, I.M. Brodo 4046 (MSC). MISSISSIPPI. Marshall Co., Holly Springs, G.T. Johnson 1030a (NY). MISSOURI. Newton Co., G. Washington Carver Nat. Mon., C.M. Wetmore 69241 (MIN); Oregon Co., Eleven Point River, H.H. Iltis (WIS). NEW JERSEY. Bergen Co., Closter, (NY); Cape May Co., Avalon, I.M. Brodo 3753 (CANL, MSC); 3757 (COLO); Lake Nummy Campsite, I.M. Brodo 3723 (MSC); Cumberland Co., E Mauricetown, J.C. Lendemer 639; Newport, J.C. Lendemer 778 (all PH); Ocean Co., Island Beach, 1942, G.G. Nearing (NY); Stafford Forge, 2000, E. Lay (personal herb.). NEW YORK. Suffolk Co., Amagansett, I.M. Brodo 3268 (MSC); Center Moriches, I.M. Brodo 2640 (MSC); Fisher's Island, I.M. Brodo 3416 (MSC, US); Gardiner' Island, I.M. Brodo 2365a (DUKE, MSC); Hampton Bays, I.M. Brodo 2296; Hither Hills State Park, I.M. Brodo 1843; Montauk, I.M. Brodo 3088 (all MSC); 3313 (UPS); Montauk Point, H.A. Imshaug 25756 (MSC); 25748 (CANL); Northville, I.M. Brodo 777; Patchogue, I.M. Brodo 623; Remsenburg, I.M. Brodo 2612 (all MSC); Shirley, I.M. Brodo 932 (COLO, MSC). NORTH CAROLINA. Durham Co., Durham, T. Tønsberg 36366 (BG). RHODE ISLAND. Washington Co., Marion Eppley Wildlife Preserve, E. Lay 06-0215 (personal herb.). SOUTH CAROLINA. Aiken Co., Aiken, H.W. Ravenel (FH, US); Berkeley Co., Santee Canal, H.W. Ravenel 188 (FH). TENNESSEE. Henderson Co., Natchez Trace, 1976, R.C. Harris (MICH). TEXAS. Cameron Co., La Fiera, A.W. Lievens 5588 (SBBG); Hidalgo Co., S Alamo, A.W. Lievens 5259 (SBBG); Houston Co, Houston, 1872, E. Hall (FH).

Reference. Brodo (1968 as R. pachysperma), Harris (1990).

TYPE. ARGENTINA. 1889, Lorentz s.n. (G, holotype)

Description. Lichenized fungus.

Thallus crustose, light gray to dark greenish gray, discontinuous or continuous becoming rimose or rimose-areolate; vegetative diaspores absent. Photobiont chlorococcoid alga, cells 8-13 μm. Ascomata lecanorine apothecia, immersed to sessile, frequent, scattered, 0.4-1.0 mm diam. Disk flat to slightly convex, dark brown to black; thalline margin sometimes poorly developed; exciple light brown, 10-30 μm wide; epithecium light to red-brown; hymenium hyaline, 60-120 μm high, not inspersed with oil droplets; hypothecium hyaline, tapering into a stipe. Asci globose clavate, 45-75 x 14-30 μm, 8-spored. Ascospores hyaline to brown, broadly ellipsoid, 2-celled, Pachysporaria-type II, 12-23.5 x 6.5-10.0 μm. Pycnidia reddish-brown, often half-exposed; conidia bacilliform, 3.0-4.5 x 1.0-1.5 μm.

Chemistry. Spot tests all negative, secondary metabolites not detected.

Substrate and Habitat. On hardwood twigs and branches, in hardwood and mixed forests.

Distribution. Eastern North America; also central and South America; in North Carolina occurs in the coastal plain and piedmont ecoregions.

Literature

Müller, J. (1889) Observationes in Lichenes Argentinenses a Doctt. Lorentz et Hieronymo lectos et a Dr. A de Krempelhubero elaboratos. Flora (Regensburg). 72: 62-68.

Sheard, J.W. (2010) The Lichen Genus Rinodina (Lecanoromycetidae, Physciaceae) in North America, North of Mexico. National Research Council of Canada, NRC Research Press, Ottawa. 246 pp.