Emmanuelia ravenelii (Tuck.) Ant. Simon & Goffinet,comb. nov. (Fig. 5E–F)

MycoBank number: MB 834654; Index Fungorum number: IF 834654; Facesoffungi number: FoF 13344;

Basionym: Sticta ravenelii Tuck., Amer. J. Sci. Arts, Ser. 2, 28: 203. 1859.

Type: USA, South Carolina: without locality; Ravenel s.n. (FH – lectotype!, here selected, MBT391294; US-US00432748 – isolectotype!). Cuba, without locality; Wright 66 p.p. (US- US00433359 p.p. – paratype!).

≡ Ricasolia ravenelii (Tuck.) Nyl., Acta Soc. Sci. Fenn. 7(2): 438. 1863.

≡ Lobaria ravenelii (Tuck.) Yoshim., J. Hattori Bot. Lab. 34: 320. 1971.

Description. Primary photobiont a trebouxioid alga. Thallus irregular in outline, small to medium to 11 cm diam., of radiating, stiff, repeatedly branched and spreading lobes. Lobes adjacent to imbricate and overlapping, to 5 mm wide between main dichotomies, with truncate to spatulate apices, adnate, with free, slightly broadly involute margins; margins entire, occasionally with small simple lobules inward. Upper surface smooth to scrobiculate, glabrous, light greenish or rarely brownish grey and light brownish toward margin when dry, greenish when wet, matte; margin arachnoid-pruinose. Lobules generally absent, rarely marginal in older parts. Medulla rather compact, white, KC+ pink. Cephalodia (with Nostoc) internal, globose to flattened, to 0.4 mm wide, widely scattered. Lower surface smooth at margin, rugose, verrucose inward, thinly tomentose, with hairs in short fascicules, light brown-cream-colored. Rhizines simple, flaring, evenly scattered, to 1.5 mm long, brown, and darkening inward. Apothecia frequent but occasionally absent, laminal, scattered to abundant, to 5 mm in diam., margin shallowly to more rarely deeply crenulated, marginal cortex pruinose, and ± rough. Upper cortex paraplectenchymatous, 20–30 µm thick, homogeneous, consisting of 4–6 cell layers. Photobiont layer 15–30 µm thick, its cells ~6 µm diam. Medulla 60–100(120) µm thick. Lower cortex paraplectenchymatous, 15–20 µm thick, with 3 cell layers; surface papillose to microtomentose in section in between short fascicles. Apothecia hymenium to 135 µm tall. Algal layer extending to the margin of apothecium. Ascospores acicular, straight to sigmoid, 1(–3)-septate, 60–75 × 3 µm, ~20–25 times as long as broad.

Secondary chemistry. Gyrophoric acid and 4-O-methylgyrophoric acid.

Ecology and distribution. On hardwood trees in the Atlantic–Gulf Coastal Plain in North America, also known from the Antilles and reported from Panama (Büdel et al. 2000).

Comments. While E. ravenelii has been considered a synonym of E. erosa [Sticta erosa] by Tuckerman (1882) and other later authors, the two species can be clearly distinguished by some morphological features, such as the absence of pruina on the upper, more distinctly scrobiculate surface of the latter, as well as a different geographical distribution (North America and Greater Antilles versus South America, respectively). Emmanuelia (as Lobaria) ravenelii has been repeatedly reported from Brazil (Kalb 1983; Brako et al. 1985). The specimens distributed in Kalb’s Lichenes Neotropici (No. 237) correspond to typical E. erosa and likely to other collections reported under this name from South America.

Specimens examined. DOMINICAN REPUBLIC. La Vega: 8.8 km N of Constanza, 6 km W of La Culata toward La Cienaga de Bermudez; 5000 ft; pine woods along road and cut-over pines with Pteridium and shrubs and a few isolated pines; 28 April 1982, R. R. Harris (B, NY). USA. Alabama, Clebourne Co., Talladega National Forest, Shoal Creek, 33.733°N 85.560°W, alt. 400 m, forested hills, on hardwood bark, 2017 December 26, V. Charny 20171226-18 (CONN); Georgia, Candler Co., Charles Harold TNC Preserve, 0–0.25 mi N of Salem Church Rd., W side of Stocking Head Creek, 32.4169°, −82.0692°, bottomland mixed hardwood forest (Nyssa, Acer, Quercus) with pine (Pinus), on Acer, 2009 December 22, J. C. Lendemer 21774 (NY #1104321); Fifteenmile Creek Preserve, E side of Fifteenmile Creek just S of I-16, 32.3614°, −82.0325°, mesic hardwood forest (Ilex, Nyssa, Fagus, Acer, Quercus) with pine (Pinus) and cypress (Taxodium) on bluff, on Magnolia virginiana, 2009 December 22, J. C. Lendemer 21699 (NY #1149787); Effingham Co., Craig Barrow farm, 32.3526°, −81.4804°, north-aspect sand hill bluff, on bark, 2012 March 4, M. F. Hodges 8253 (NY #2057430); Emanuel Co., Ohoopee Dunes, 32.5253°, −82.4472°, on bark, 2007 October 7, M. F. Hodges 1941-C (NY #2057238); Tattnall Co., Big Hammock Natural Area, blue and yellow trails, ~1 mi E of jct of CR 441 & GA 121/144/169, 31.8576°, −82.0589°, sand hill community with oak scrub and mixed hardwoods, on Carya, 2009 December 20, J. C. Lendemer 21443 (NY #1886944); Big Hammock Natural Area, ~1 mile southeast of Highway 121/144/169 off of Mack Phillips Road., 31.8636°, −82.0525°, alt. 20 m, dense forest with Pinus palustris, Quercus spp., and Prunus sp., on bark, 2013 March 25, T. S. Quedensley 10852 (NY #1150285); Mississippi, Wilkinson Co., 15 min. NW of St Francisville, between road 969 and the river, deep ravines with hardwood forest, on a deciduous tree in the bottom of the ravine, May 29 1976, E. Sérusiaux 1666 (LG); North Carolina, Craven Co., Croatan National Forest, 0–0.2 mi NE of FS 170, along tributary to Brices Creek, 1.9 mi NE of jct of FS170/FS121.2 and FS1101, 35.0369°, −77.0497°, alt. 4 m, swamp forest of mixed hardwoods (Ilex, Acer, Nyssa, Liquidambar, Persea, Magnolia virginiana), on Acer, 2013 March 5, J. C. Lendemer 35039 (NY #1865943); Dare Co., Alligator River National Wildlife Refuge, Brier Hall Rd. 0.25–0.5 mi S of jct w/ Mallard Rd., 0.25 mi N of US64, 35.8828°, −75.9375°, alt. 1 m, mixed hardwood (Ilex opaca, Acer, Liquidambar, Nyssa, Persea) – Pinus swamp with understory of Cyrilla and Ilex glabra, on Acer, 2014 March 21, E. A.  Tripp 4654 (NY #2203723); Dare Co., Alligator River National Wildlife Refuge, W of Whipping Creek Rd. 0.5 mi N of jct w/ Chip Rd., 35.6753°, −75.9625°, alt. 0 m, mature mixed hardwood (Nyssa, Acer, Magnolia virginiana, Liquid- ambar) – conifer (Chamaecyparis [dead], Taxodium and Pinus) swamp forest with Ilex glabra and Clethra understory, on large Acer, 2014 March 23, J. C. Lendemer 43114 (NY #2203643); Graham Co., Nantahala National Forest, W of Powell Branch at confluence with Fontana Lake, terminus of Cable Cove Road (CR 1287) at Cable Cove Boat Launch, 35.4372°, −83.7481°, alt. 527, stream ravine with acid hardwood-conifer forest, on poison ivy, 2016 March 18, R. C. Harris 61074 (NY #2467099); Washington Co., Bull Neck Swamp, Deep Creek Rd. 0.5 mi N of bridge over Deep Creek, 35.9372°, −76.3992°, alt. 0 m, mixed hardwood (Nyssa, Acer, Magnolia virginiana)-Taxodium swamp forest, on base of Nyssa, 2014 March 22, W. R. Buck 63035 (NY #2327942); South Carolina, McCormick Co., Sumter National Forest, Stevens Creek Heritage Preserve, N of SSR 88, ~1.5 mi NE of Clarks Hill, 33.6872°, −82.1614°, alt. 100 m, mesic mixed hardwood bluff forest, on rock, 2010 March 11, R. C. Harris 55880 (NY #1148729).

Molecular data. Yes.

Figure 5. Habit of Emmanuelia species. A – E. americana (Lücking 40112), showing upper side with pycnidial warts; B – E. erosa (Cáceres 25148), showing upper side with distinct scrobiculate surface; C–D – E. elaeodes (C, Lücking 40082; D, Lücking 37546), showing upper side with apothecia; E–F – E. ravenelii (Hollinger 2332, CONN), showing upper side with numerous apothecia (E) and close-up of lobe margins with pruinose surface (F). Photographs E & F: courtesy of Jason Hollinger.