Taxonomic Note. The description below, copied from Sheard (2010), refers to R. destituta, which, according to Sheard (2018), is a synonym of R. moziana.

Basionym. Lecidea destituta Nyl., Sertum Lich. Trop. Labuan et Singapore, Accedunt Observ.: 41 (1891).
Type. U.S.A. ILLINOIS. Super saxa arenaria, s.d. J. Wolf s.n. (H‑NYL 10210 ‑ holotype!).

Rinodina ochrocea Willey ex Hedrick, Mycologia 26: 105 (1934).
Type. U.S.A. SOUTH CAROLINA. Chester Co., on rocks, s.d. H.A. Green s.n. (MICH ‑ holotype!, US - isotype!).

Rinodina vezdae H. Mayrhofer, J. Hattori Bot. Lab. No. 55: 473.
Type. Vězda Lich. Bohem. exs. 113 (as R. discolor), „SSR. Moravia occid.: Vev. Bitýška, ad saxa dioritica in valle fluvii Svratka, ca. 350 m, 12. 12. 1956, A. Vězda s.n. (M - holotype!).

Exsiccatae. Vězda Lich. Bohem. Exs. 113 (M as R. vezdae).

Description. Thallus thick, light to dark grey, areolate, rarely rimose, rarely inconspicuous and chasmolithic; areoles typically isolated, verrucose, to 0.30-0.50 mm wide at first, coalescing to form areoles 1.20-1.60 mm wide, sometimes with subsquamulose margins; surface rugose, rarely plane, matt; margin indeterminate; prothallus lacking or rarely fimbriate; vegetative propagules absent. Apothecia innate, becoming broadly attached, frequent, often contiguous, to 0.50-0.90 mm in diam.; disc dark brown (particularly when wet) to typically black, persistently plane; thalline margin concolourous with thallus, entire, sometimes incompletely formed or becoming pigmented in part, persistent, 0.05-0.10 mm wide, or rarely absent; excipular ring sometimes present, slightly raised. Apothecial Anatomy. Thalline exciple rarely absent, typically 60-100 µm wide laterally; cortex 10-15 µm wide, outer ca. 25 µm sometimes becoming pigmented, dark brown or aeruginose (Bagliettoana-green, N+, K+ red); epinecral layer absent; crystals present in cortex, absent in medulla; cortical cells to 4.0-5.0(-6.0) µm wide, not pigmented; algal cells to 9.0-17.0 µm long; thalline exciple (60-)80-110 µm wide below; cortex ca. 10 µm wide; proper exciple hyaline, 5-10 µm wide laterally, to 10-30 µm wide above, sometimes aeruginose above, or 40-60 µm wide when thalline exciple absent; hypothecium 50-70(-90) µm deep; hymenium 90-110 µm high, not inspersed; paraphyses 2.0-3.0 µm wide, typically conglutinate, with apices to 3.0-4.5(-5.5) µm wide, brown capitate forming a light brown epihymenium; asci 60-90 x 17-29 µm. Ascospores 8/ascus, Type A development, Mischoblastia-type, (19.0-)22.0-22.5(-26.0) x (9.5-)12.0-13.0(-15.0) µm, l/b ratio 1.7-1.9, largest spores often swollen at septum, lumina sometimes becoming inflated but retaining thick apical wall at maturity; torus narrow at maturity; walls lightly pigmented, not ornamented. Pycnidia immersed in thallus, ostiole pigmented, < 0.05 mm wide; conidiophores type I; conidia bacilliform, 3.0-4.5 x ca. 1.0 µm.

Chemistry. Spot tests, K+ yellow, C-, KC-, P+ faint yellow; secondary metabolites, atranorin in cortex.

Substrate and Ecology. Saxicolous, frequently on granite and other siliceous rocks, rarely on limestone, in moist, open and glade habitats.

Distribution. The species is frequent throughout the Great Lakes region and common in southern Ontario and Minnesota. Further south it has a more scattered distribution. Recorded as R. vezdae by Lendemer and Macklin (2006), it belongs to the Atlantic element and approximates the Appalachian Province of Flora North America (1993). The species is otherwise found in central and southern Europe, including Spain where it is rare (Giralt 2001).

Notes. Rinodina destituta is related to R. oxydata in possessing Mischoblastia-type spores, an apothecial margin which may become pigmented, and by the presence of atranorin in the cortex. It is separated from the latter species by the thicker thallus, typically with relatively large, rugose to verrucose areoles, and spores averaging >21 µm long. Young thalli are characterized by thin, convex and noncontiguous areoles, and therefore distinguished from thalli of R. oxydata which possess plane, contiguous areoles in early stages of development. Nevertheless, young thalli of R. destituta are not always easy to distinguish from R. oxydata because their spores may also be smaller than typical. Thicker thalli of R. oxydata are distinguished by their plane rather than rugose surface.

There is a considerable range of thallus colour and morphology, most variation apparently being due to shade and moisture levels. Shade results in darker thalli and often a reduction in atranorin content as deduced from its crystal concentration in the cortex when viewed under polarized light. Increasing moisture results in the thallus surface becoming less rugose and the areole margins being raised from the substrate, this effect sometimes being visible within a given thallus where it occupies moisture retaining hollows in the substrate. There may also be a trend for this morphotype to be more common in the southern part of the species range. The holotype of R. ochrocea belongs to this morphotype with the additional character of its ochraceous colour, probably the result of inundation by iron stained water. A second, larger specimen from the type locality in MICH, dated 10-III-1888, is probably an isotype. Previously known as R. vezdae in Europe, the holotype of that species is intermediate in morphology between the typical verrucose form of the Great Lakes region and the “ochrocea” morphotype.

Another extreme morphotype is represented by the type of R. destituta. The thallus of this morph is very reduced being limited to cracks between quartz crystals (chasmolithic) of its coarse sandstone substrate. The other characteristic of this morph is the lack of algae and hyaline medullary tissue in the margin which is otherwise of rare occurrence in the typical form with a well developed thallus. The spores are almost identical in size to the typical form but the apothecia are slightly smaller which may be consistent with the poorly developed thallus. Algae are present in the usual abundance below the hypothecium, only rarely extending slightly into the base of the margin. Atranorin has not been confirmed in the type but the presence of crystals under polarizing light suggests that it is present. This morphotype represents an extreme form of the species related to its atypical substrate.

Clearly there is a great deal of variability within this species as there is within the related R. oxydata. For these reasons Giralt et al. (1997) included it (as R. vezdae) within R. oxydata but declined to formally place it into synonymy. Collections of this species were named R. lecanorina by Fink (1910, Fink 901 Koochiching (MU), Fink 1072 Rainy Lake City (MICH)) and subsequently R. ocellata (Fink 1935). The species is referred to as R. “ destitula” in Fink (1935). Specimens reported as R. iowensis by Brodo (1988) and R. verrucosa by Brodo (1981) also belong to R. destituta.

Specimens from California, growing on soil and loosely compacted conglomerate, possess significantly larger spores (averaging 25.5-27.0 x 13.0-14.0 µm), with a well developed torus and more strongly ornamented walls, probably belong to a separate species. I have declined to recognize this material formally as the few specimens studied are in poor condition due to the nature of the substrate and because the whole group of species with Mischoblastia-type spores is in need of revision (Matzer and Mayrhofer 1996). Only one recent collection has been seen (Knudsen 6203, from Orange Co., California, SASK, UCR). Such material was included by Hasse (1913) under R. conradii, R. turfacea (Hasse Lich. exs. 249, SBBG) and R. turfacea var. mniaraea.

Another potential taxon exists in Pudget Sound-Straits of Georgia area of the Pacific Northwest. This population is characterized by larger spores, smaller conidia and algal cells, and with the Bagliettoana-green pigment of the upper proper exciple and adjacent thalline exciple sometimes extending into the epihymenium in low concentrations. The British Columbia specimens were referred to as R. verrucosa Sheard ined. by Noble (1982). This name was first used as a herbarium name for material now included in R. destituta and was published by Wong and Brodo (1973) for specimens recorded from Ontario.

Species with Mischoblastia-type spores are all saxicolous except for R. euskadienis (Crespo and Aguirre1984) which was described growing on Fagus sylvatica in Spain. Giralt (2001) compares this species to R. oxydata because of its atranorin containing cortex but its large spores suggest it might be closer to R. destituta.

Specimens examined. CANADA. MANITOBA. Winnipeg, 1927, K. Scott (MICH). NEW BRUNSWICK. Point Wolfe, S. Gowan 2456 (CANL). ONTARIO. Thunder Bay Dist., Little Dog Lake, R.F. Cain 20257 (UPS); Carleton Co., Bells Corners, P.Y. Wong 1863; Brittania, 1895, J. Macoun; Hogs Back, J. Macoun 1905; Rockcliffe, 1897, J. Macoun; Rockcliffe Park, I.M. Brodo 22776a (all CANL); Ottawa, R.E. Lee 1185, 1339 (personal herb.); South March, M.J. Shchepanek 9; Frontenac Co., Clarendon, P.Y. Wong 2899; Lake Opinicon, P.Y. Wong 302, 478; Glengarry Co., Alexandria, P.Y. Wong 1197; Grey Co., P.Y. Wong 1940; Hastings Co., Marmora, P.Y. Wong 2258 (all CANL); Lanark Co., Indian Creek, R.E. Lee 1251 (personal herb.); Maberley, P.Y. Wong 3204 (CANL); White Lake, R.E. Lee 1289 (personal herb.); Leeds Co., Forthton, P.Y. Wong 2279; Lennox & Addington Co., Camden East, P.Y. Wong 2472; Peterborough Co., Stoney Lake, P.Y. Wong 4416; Renfrew Co., Eganville, P.Y. Wong 3027; 6 mi N Eganville, I.M. Brodo 20570B; La Passe, P.Y. Wong 3013 (all CANL). U.S.A. ALABAMA. T.M. Peters (FH). DELAWARE. New Castle Co., Faulkland, 1886, A. Commons; Newark, 1892, A. Commons (both PH). ILLINOIS. Rock Castle Creek, A.C. Skorepa 6232 (BALT); Calhoun Co., Cap au Gres, M.D. Jones 2306; Grundy Co., Goose Lake Prairie, G. Wilhelm 19266; Hardin Co., Finneyville, M.A. Basinger 9031; Kendall Co., 4 mi S Oswego, G. Wilhelm 13953; La Salle Co., Sheriden Red Pine, M.D. Jones 1627b; McHenry Co., McHenry, G. Wilhelm 19323 (all MOR); Randolph Co., Piney Creek Ravine, A.C. Skorepa 6221, 6295 (BALT). IOWA. Butler Co., Seven Mile, 1909, B. Fink (MICH). KANSAS. Douglas Co., Baldwin, C.L. Kramer 495 (WIS); Ellsworth Co., 4mi SE Carneiro, C.A. Morse 12700; 2mi SW Carneiro, C.A. Morse 12652; Montgomery Co., 5.5m1 W Liberty, C.A. Morse 14379b; Russell Co., 7.5mi S Lucas, C.A. Morse 12293; Russell Co., 7.5mi S Lucas, C.A. Morse 12293 (all KANU); Wilson Co., 11 mi NW Fredonia, R.A. Anderson 29099 (COLO); Woodson Co., 1mi W Rose, C.A. Morse 14487 (KANU).  LOUISIANA. Union Parish, 6 mi S Bernice, J. Kay 8839 (MSC). MARYLAND. Soldier's Delight, 1911, C.C. Plitt (FH, US); Baltimore Co., 1912, C.C. Plitt (US); Bare Hills, 1911, C.C. Plitt (FH, US); Cecil Co., S Porters Bridge, 2002, J.C. Lendemer (PH); Montgomery Co., Plummer's Island, B. Fink (FH); B. Fink 60 (US); 1908, B. Fink (MU, UPS, US). MASSACHUSETTS. Hampshire Co., Amherst, 1861, E. Tuckerman (FH). MICHIGAN. Ontonagon Co., Porcupine Mountains, 1964, R.C. Harris (MSC). MINNESOTA. Big Stone Co., 2 mi SE Ortonville, G.A. Wheeler 18510 (MIN); Blue Earth Co., Mankato, B. Fink 127 (MICH, MIN); Chisago Co., St Croix River, C.M. Wetmore 62796 (MIN); Koochiching Co., Koochiching, B. Fink 901 (MU); B. Fink 981 (MIN); Rainy Lake City, B. Fink 1072 (MICH); B. Fink 1081; Lac qui Parle Co., 6 mi NNE Bellingham, G.A. Wheeler 18536; Lake Co., SW Sourdough Lake, T.D. Trana 13857; Lake of the Woods Co., Oak Island, B. Fink 511; Nicollet Co., 2.5 mi NW Courtland, G.A. Wheeler 17460; 4.5 SE Fort Ridgely State Park, G.A. Wheeler 18722 (all MIN); Pine Co., Kingsdale, J.P. Schuster 1491 (SASK); Redwood Co., 5.5 mi N Belview, G.A. Wheeler 16459; Renville Co., Morton, B. Fink 316; Scott Co., 4.5 NNE Jordan, G.A. Wheeler 19029; St. Louis Co., Voyageurs Nat. Park, C.M. Wetmore 31867; Stearns Co., 2 mi NW Rockville, G.A. Wheeler 16645; 3.9 mi S St. Stephens, G.A. Wheeler 16715; Winona Co., 20 mi NW Winona, L. Brako 1291; Yellow Medicine Co., 5 mi NE Echo, G.A. Wheeler 16490 (all MIN). MISSOURI. E. Hall 24 (FH); Carroll Co., Bunch Hollow, G. Wilhelm 16050; Cranford Co., Onondaga Cave State Park, G. Wilhelm 20031 (both MOR). NEBRASKA. Lancaster Co., 1899, Sheldon (FH); Lincoln, 1936, W. Kiener (NY). NEW YORK. Erie Co., Buffalo (FH); Madison Co., Bridgeport, 1958, I.M. Brodo (CANL). OHIO. Butler Co., Darrtown, 1909, B. Fink (MICH); Near Oxford, 1927, B. Fink (WIS); Oxford, 1927, B. Fink (MICH); Reily, 1909, B. Fink  (MICH); Seven Mile, 1909, B. Fink (MICH, WIS); Delaware Co., 1935, J. Wolfe 20 (CANL); Fairfield Co., Amanda, 1892, W.S. Kellerman (MU); Franklin Co., Columbus, 1892, E.E. Bogue (FH); Geauga Co., Bainbridge, B. Fink 437; Preble Co., Eaton, 1914, B. Fink 200, 205, 229; West Alexandria, B. Fink 318; Warren Co., Mason, B. Fink 523 (all MICH). OKLAHOMA. Comanche Co., Beaver Bend Park, J.W. Thomson 8670; Payne Co., 6 mi W Stillwater, J.W. Thomson 8956 (both WIS). SOUTH CAROLINA. Chester Co., H.A. Green (MICH); 1885, H.A. Green (FH); 1886, H.A. Green (US); 1888, H.A. Green (MICH); 1896, H.A. Green (NY); Landsford, 1884, H.A. Green (PH). TEXAS. Burnet Co., 2 km S Inks Lake, R.S. Egan 9191 (MIN). VERMONT. Windham Co., Battleboro, C.C. Frost 44 (FH). VIRGINIA. Highland Co., 9 mi S Monterey, R.C. Harris 9959, 10687 (MICH). WISCONSIN. Trempealeau Co., Ettrick, K.G. Foote 1269 (WIS); Hollandale, 1925, G.K. Merrill (FH); Ashland Co., Copper Falls State Park, I.M. Brodo 5680b (CANL, MSC); Douglas Co., Superior, J.W. Thomson 2155 (WIS, UPS); 2165 (UPS); Polk Co., Nevers Dam Landing, C.M. Wetmore 67812; Sunrise Landing, C.M. Wetmore 67786 (both MIN).

Selected References. Wong & Brodo (1973), Brodo (1981), Noble (1982) all as R. verrucosa unpublished name, Mayrhofer (1984a as R. vezdae), Brodo (1988 as R. iowensis), Giralt (2001 Plate VII: B as R. vezdae), Thomson (2003 partly as R. confragosa).

Sheard, J. W.  (2018)  A synopsis and new key to the species of Rinodina (Ach.) Gray (Physciaceae, lichenized Ascomycetes) presently recognized in North America.  Herzogia 31(1): 395-423.