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Stretching the Boundaries: A Range Extension for Buellia wheeleri R.C. Harris JAMES C. LENDEMER Cryptogamic Herbarium, Institute of Systematic Botany, The New York Botanical Garden, Bronx, NY, 104585126, U.S.A.; e-mail: [email protected]

BRENDAN P. HODKINSON Department of Biology, Duke University, Durham, NC, 27708, U.S.A.; e-mail: [email protected]

Abstract. - Buellia wheeleri is newly reported from a number of locations throughout the southeastern Coastal Plain of North America (e.g., Alabama, Arkansas, Georgia, Tennessee, Virginia, and the Delmarva Peninsula of Maryland). This distinctive species was previously known in North America only from a handful of collections in Florida and North Carolina.

Recently, the second author spent nearly a week at The New York Botanical Garden (NY) working on the identification of problem specimens collected during his fieldwork (Hodkinson et al. 2009; Hodkinson in press). Among these specimens was a sterile sorediate crustose lichen on oak bark, collected in eastern Virginia. Initially, we were unable to identify the specimen, but the first author recognized it as the same taxon he had collected on the Delmarva Peninsula in Maryland several years earlier and bore a resemblance to Biatora pontica Printzen (Lendemer & Knapp 2007). The two collections were highly distinctive because of their C+ and KC+ yellow-orange spot tests and punctiform soralia with intense yellow soredia. The week prior to the second author’s visit to NY, we had both attended the Tuckerman Workshop which was held in southern Florida. During the meeting, we

made several collections of Buellia wheeleri R.C. Harris (treated in the monotypic genus Ciposia Marbach as C. wheeleri (R.C. Harris) Marbach by Marbach (2000)), a sorediate crustose species that is frequently sterile. While processing these collections upon our return home, we realized that our specimens from Maryland and Virginia represented B. wheeleri. Buellia wheeleri was originally described by Harris (1988) from Polk County, Florida, and was subsequently found to be common on the bark of oaks throughout the central and northern portions of the state (Harris 1995). It was soon reported from southern Illinois, though at the time this would have seemed to be a rather anomalous report for a potential Florida endemic (Wilson and Methven 1997); unfortunately, a voucher for this report could not be located. Lendemer and Yahr (2004) later reported it

173 (again on the bark of oak trees) from coastal North Carolina. In light of these reports, one would expect B. wheeleri to have a wider distribution in the southern coastal plain of southeastern North America, comparable to other species that occur on substrates (e.g., Taxodium) or in habitats (e.g., oak-scrub) that are common throughout this phytogeographic province. Species that display this type of distribution are typically common in central-northern Florida and become rarer, eventually disappearing, as one moves further north along the Atlantic coast (see discussion in Lendemer & Knapp (2007) and Lendemer (2006)). Unfortunately, our knowledge of the distributions of many taxa in this region, especially asexually reproducing crustose lichens, is hindered by a lack of reliable reports and adequate modern collections. A distribution map for Buellia wheeleri is presented here (Fig. 1) with a list of selected specimens examined. In addition to the records mentioned above, the herbarium of The New York Botanical Garden (NY) also contains material of B. wheeleri from the far reaches of the coastal plain in Arkansas and Tennessee. The species has also recently been collected in Georgia. While the majority of collections occur on Quercus, there are also specimens from Acer, Sabal, and Taxodium. As noted above, Buellia wheeleri is a frequently sterile sorediate crustose lichen that typically occurs on the rough bark of the trunks of oaks (Quercus) in oak-scrub habitats. The thallus is dark green and comprised of thin continuous to dispersed areoles, although in some populations these areoles are poorly developed or even absent (see Figs. 2a & 2b). Since there are few

Volume 26 (4) sorediate species of Buellia, the combination of soredia and large, strongly ornamented ascospores (16-19 x 9-11.5 µm fide Harris (1995); see illustrations in Harris (1988)) serve to distinguish B. wheeleri from any other North American species currently recognized in the genus. Apothecia and ascospores are often immature or entirely absent, and so spores are a poor diagnostic character for the majority of specimens. However, specimens can be recognized in the field, even when sterile, because of their substrate preference, small punctiform to diffuse, confluent soralia, and soredia that are intensely yellow due to the presence of the pigment secalonic acid (K-, C+, KC+ yellow-orange). Without spot test data, specimens representing this taxon are most likely to be confused with sterile species of Caloplaca (K+ purple) or Candelariella (C-, KC-). In eastern North America, only Biatora pontica might be confused with Buellia wheeleri; however, that species occurs in the Appalachian Mountains, not the coastal plain, and owes its color to the presence of a xanthone instead of secalonic acid. The present note represents the first attempt to characterize the distribution of Buellia wheeleri in North America since its description in 1988, and we anticipate that future collections will fill in the distribution of this distinctive southeastern Coastal Plain species. Selected Specimens Examined. (all on Quercus unless otherwise indicated). U.S.A. ALABAMA. BALDWIN CO.: Shelby’s, 8.iv.1895, on palmetto, A.W. Evans 429 (NY). ARKANSAS. JEFFERSON CO.: Pine Bluff Arsenal, W of N end of Yellow Lake, 2.xii.1999, on

174 Taxodium distichum, D. Ladd 21972 & M. Pederson (NY). FLORIDA. BAKER CO.: along CR 127 at Moccasin Creek, 26.xi.1996, R.C. Harris 39295 (NY). CALHOUN CO.: along S side of FL 20, ~8 mi E Bay Co. line, 30.xi.1988, W.R. Buck 16505 (NY). COLUMBIA CO.: Osceola National Forest, W of FS 237, 15.xii.1993, R.C. Harris 32540 (NY). DIXIE CO.: Steinhatchee Wildlife Management Area, 4.xii.1993, R.C. Harris 31629 (NY). HIGHLANDS CO.: Hickory Hammock, 28.iii.1998, R.C. Harris 41905 (NY). LEON CO.: Apalachicola National Forest, Leon Sinks Geological Area, 2.xii.1988, R.C. Harris 23279 (NY). OKEECHOBEE CO.: Viking Property, 28.iii.1998, R.C. Harris 41881 (NY). ORANGE CO.: along CR 532, 2 mi SSW of SR 520, 7.i.1996, R.C. Harris 37489 (NY). POLK CO.: Ridge Audubon Center, 16.i.1989, E.M. Wheeler s.n. (NY). PUTNAM CO.: Ocala National Forest, old Johnson Field Campground, 7.xii.1988, W.R. Buck 16815A (NY). SANTA ROSA CO.: Blackwater River State Forest, 9.xii.1993, R.C. Harris 32042 (NY). SARASOTA CO.: Longboat Key, 26.ii.1937, C.S. Romer s.n. (NY). SEMINOLE CO.: along Willingham Rd., 0.5 mi N Co. Rd. 419, 10.i.1996, R.C. Harris 37761 (NY). ST. JOHNS CO.: South Ponte Vedra Beach, 28.xi.2008, W.R. Buck 54360 (NY). TAYLOR CO.: Big Bend Wildlife Management Area, 3.xii.1996, R.C. Harris 39446 (NY). UNION CO.: Lake Butler Wildlife Management Area, 4.xii.1994, R.C. Harris 35998 (NY). GEORGIA. GLYNN CO.: Jekyll Island, 8.iii.2009, S.Q. Beeching 7936 (GAM). MARYLAND. CAROLINE CO.: Idylwild State Wildlife Management Area, 18.iv.2006, J.C.

Volume 26 (4) Lendemer 6227 & W. Knapp (NY). NORTH CAROLINA. BRUNSWICK CO.: Bald Head Island, 5.vii.2003, J.C. Lendemer 1321 & R. Yahr (NY). CARTERET CO.: Cape Lookout National Seashore, Shackleford Banks, 19.iii.2003, R.C. Harris 47198 (NY); Bogue Banks, 20.iii.2003, W.R. Buck 43863 (NY). TENNESSEE. HENDERSON CO.: Natchez Trace State Park, 4.vii.1976, R.C. Harris 11571 (NY). VIRGINIA. HENRICO CO.: Cheswick Park, 02.viii.2008, B.P. Hodkinson 10239 (NY); Woodman Road NWF Wildlife Habitat, 05.iv.2009, B.P. Hodkinson 10939 (DUKE).

ACKNOWLEDGMENTS Many thanks to Andrew Methven (EIU) for help locating specimens associated with this and other related projects. Also thanks to Robert Lücking (F) and the National Science Foundation for providing funding that allowed both authors to attend the 2009 Tuckerman Workshop (NSF Award #: DEB-0715660). We are grateful to Sean Beeching for sharing his observations and unpublished reports of B. wheeleri in Georgia, and to Richard C. Harris for reviewing this manuscript. Finally, we thank Donald and Beverly Ziegler for providing field assistance and access to their Wilderness Area in Henrico, Virginia. LITERATURE CITED Harris, R.C. 1988. Buellia in North and Central Florida or the virtues and rewards of collecting. Evansia 5(3): 3745. Harris, R.C. 1995. More Florida Lichens. Including the 10¢ Tour of the Pyrenolichens. Publ. by the Author, Bronx, N.Y. 192 pp.

175 Hodkinson, B. P., R. C. Harris, and M. A. Case. 2009. A Checklist of Virginia Lichens. Evansia 26(2): 64-88. Hodkinson, B. P. 2010. A First Assessment of Lichen Diversity in One of North America's 'Biodiversity Hotspots' in the Mountains of Virginia. Castanea 75: in press. Lendemer, J.C. 2006. Contributions to the Lichen Flora of New Jersey: A Checklist of the Lichens of Wharton State Forest. Opuscula Philolichenum, 3: 21-40. Lendemer, J.C. and W. Knapp. 2007. Contributions to the Lichen Flora of Maryland: Recent Collections form the Delmarva Peninsula. Opuscula Philolichenum 4: 21-40.

Volume 26 (4) Lendemer, J.C. and Yahr, R. 2004. A checklist of the lichens collected during the Tuckerman Workshop #12, Outer Banks, North Carolina, USA. Evansia 21(3): 118-136. Marbach, B. 2000. Corticole und lignicole Arten der Flechtengattung Buellia sensu lato in den Subtropen und Tropen. Bibliotheca Lichenologica 74, J. Cramer, Berlin, Stuttgart. 384 pp. Wilson, P. J. and Methven, A. S. 1997 (1998): Lichen use by larval Leucochrysa pavida (Neuroptera: Chrysopidae). The Bryologist 100(4): 448-453.

Figure 1. Geographic distribution of Buellia wheeleri as presently known (based on specimens from DUKE, NY, and hb. Beeching). The shaded area approximates the extent of the coastal plain in North America.

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Figures 2a-b, both from Buck 54360 (NY). Thallus and apothecia of Buellia wheeleri (left, x3). Young punctiform soralia (right, x7).