Vaccinium ashei J.M.Reade, Torreya

Vaccinium ashei J.M.Reade, Torreya View in CoL 31:71.1931.

TYPE: U.S.A. FLORIDA. Okaloosa Co.: near Niceville , in small swamp with titi , Cyrilla etc., 26 Mar 1927, W.W.Ashes.n. (LECTOTYPE, designated here:GA accession #275541,barcode #259756, flowering branchlet on sheet [lower element] , Fig.12 View FIG .See also Fig. 13a–g View FIG for photographs of living plants illustrating diagnostic features of V.ashei .

Reade (1931) described the type of Vaccinium ashei as follows: “The type material collected by W. W. Ashe near Niceville, Okaloosa County, Fla., consisting of both flowers, March 26, 1927, and fruit June 2, 1929, from the same plant, is in the Ashe herbarium. Co-type specimens in fruit are being deposited in the herbarium of the University of Georgia and in the National herbarium.” The “Ashe herbarium” may be at NCU, but we have located no type material there. The only type material we have located is at GA. W. H. Camp annotated this specimen as “ ISOTYPES ” in 1942 and “ GA Herbarium Staff” annotated the specimen as isotype in 2017, but it is not clear if any of these individuals saw material that could be considered the holotype. Because the GA specimen constitutes the only type material that we have seen, and because there is no evidence of the existence of type material at the “Ashe herbarium,” whether that is NCU or a different herbarium, we have used the material on the sheet at GA for lectotypification. There are two elements on the sheet, one in flower and one in fruit, each collected at a distinct time and therefore each constituting a separate gathering; as such, these are syntypes. We designate the flowering specimen on the sheet as lectotype and presume that the date of collection is the date indicated in the protologue, although only the fruiting date of 8 June 1929 is indicated on this sheet. Reade did not annotate this specimen, but because GA is Reade’s herbarium, we presume that Reade saw it. We have not seen the specimen at US (presumably the “National herbarium” referenced by Reade).

Description.— Data from Reade (1931) that differ from our observations are indicated in brackets. Shrubs, deciduous, usually clumped and multi-stemmed, not clonal or ( Camp 1945) clonal, 1.0–4.5[–6.0] (rarely –7, from Camp 1945) m t all, top-most branches often arched, distal portions drooping in fruit. Branchlets in situ green until ca. third or fourth year, older branchlets and stems gray to brown; eruptive periderm (split longitudinally and irregularly) common on third-year and older branchlets and stems. Current-year branchlets sparsely to densely white-puberulent, without stipitate glandular trichomes, trichomes in 2 narrow lines, straight to curved, 0.10–0.34 mm long; second-year branchlets glabrous or white-puberulent, glabrate, trichomes ± in lines and/or patches, often broken; outer vegetative bud scales 1.3–2.4 mm long, glabrous or occasionally puberulent. Leaves with petiole 0.5–2.2[–3.0] mm long, sulcate and/or margins narrowly winged to base, abaxially and adaxially pubescent, eglandular; leaf blade elliptic or narrowly elliptic to ovate or slightly obovate, often ± rhombic, (2.3–)3.4–7.9 × (0.9–)1.6–3.8[–4.3] cm, coriaceous, abaxial surface in situ dull, bluish green and glaucous, in sicco often pale tan-brown or brown, distinctly paler than adaxial surface, pubescent with white ± apically-oriented ± curved trichomes 0.32–0.50(–0.60) mm long borne along midvein at least from base to distal 1/3 and occasionally secondary veins, variously becoming less pubescent with age, sparsely to less often moderately stipitate-glandular along veins and occasionally on surface (stipitate-glandular trichomes 0.12–0.38 mm long, stipes usually not notably flattened, not notably flanged proximally, gland-heads orangish, reddish, or black, globose to ovoid), adaxial surface in situ bluish green and glaucescent, in sicco bluish green to brown to dark brown and glaucescent, glabrous except for ascending to erect ± curved eglandular trichomes along midvein, base cuneate to broadly cuneate, margins occasionally entire or usually crenulate-serrulate with each tooth tipped by a stipitate gland, sparsely to densely ciliate-pubescent at least proximally, narrowly revolute, tapering into petiole, apex [obtuse to] acute to slightly acuminate, sessile marginal glands present, 0.06–0.16 mm wide (width parallel to margin), marginal teeth (not including stipitate gland) when present oriented toward leaf apex with straight to outcurved outer edges and shallow and narrow to deep and broad sinuses. Inflorescences of axillary or pseudoterminal racemes, borne at most at several contiguous nodes along non-elongate or non-virgate distal portions of branchlets, 2- to 9-[to 10-]flowered, rachis 1–11 mm long, subtending bracts white often strongly flushed pink, obovate to subrotund, cucullate, glabrous (not stipitate-glandular), margins eciliate, pedicels glabrous, bracteoles white often strongly flushed pink, narrowly elliptic to narrowly rhombic, glabrous (not stipitate-glandular), margins eciliate. Flowers: emerging before leaves and continuing nearly to full leaf size; calyx hypanthium 0.6–2.3 × 1.3–3.3 mm, glabrous, with glaucescence on at least some hypanthia or (observed in sicco, extreme eastern part of range) rarely completely without; calyx limb 0.4–1.2 mm long; calyx lobes deltoid to ± hemispheric, 0.5–1.5 × 1.4–2.1 mm, with or without glaucescence, margins eciliate or slightly ciliate, apex acute to rounded; corolla white, occasionally tinged reddish [rarely striped with red], ± cylindrical, broadest in middle third, slightly constricted subapically, 6–10 × 3.5–6.5 mm, glabrous within and without, except often with sparse to dense short trichomes on inner lobes; stamens 6.4–8.0 mm long; filaments 2.6–4.4 mm, glabrous on surface, white-hirtellous marginally, trichomes to 0.70 mm long; anthers 3.8–4.9 mm long, thecae 1.5–2.0 mm long, tubules 2.1–3.0 mm long; style exserted, 8–10 mm long, glabrous. Fruits pastel green when immature turning pinkish pale green to wholly reddish, and black or purplish black at maturity, with or without glaucescence, subglobose, 8–16 mm diam., with sweet flavor; seeds ca. (20–) 23 to 44 in number, chestnut brown, 1.1–1.7 × 0.8–1.2 mm.

Distribution habitat, and floral phenology.— The natural geographic range of Vaccinium ashei is here considered to extend from extreme southeastern Louisiana east to extreme southeastern Georgia and extreme northeastern Florida. The natural habitats occur within the longleaf pine ( Pinus palustris Mill. ) ecosystem. Specimen labels yield the following habitat descriptors, generally ranging from moist or wet creek-sides and river margins to mesic pine-hardwood forests, mature pine forests, and dry pine-oak woodlands, margin of the Okefenokee Swamp, bottomland woods by creek, sandy seepage along river, edge of hammock above creek, moist woods on slope above creek, moist Magnolia -spruce pine woods, Magnolia -pine creek bottomland, ecotone of stream head, wooded strip around pond, roadcut bank near creek, roadsides, mesic to moist longleaf pine slopes, cut-over longleaf pine, flat open pineland, open pine-live oak-laurel oak woods, well-drained mixed woods, oak woods on sandy ridge, and very sandy pine-live oak-myrtle oak woods. Flowering occurs from late February to mid-April; fruiting occurs from early June to mid-August.

Chromosome number.— 2n=6 x =72 (hexaploid).

PRESUMED ESCAPES FROM CULTIVATION OF RABBITEYE BLUEBERRY

We encountered several herbarium specimens and plants in the field (which we vouchered as herbarium specimens; e.g., Fig. 13h View FIG ) that are morphologically similar to Vaccinium ashei in most respects. In most of these specimens, however, the current-year stems are either glabrous (versus with some pubescence in V.ashei as circumscribed here) or more sparsely pubescent than those of V. ashei and the abaxial surfaces of the leaf blades are completely glabrous (versus always at least sparsely pubescent). Furthermore, they were collected either at the periphery of the range of V. ashei as here circumscribed or well outside this range. From this we conclude that these specimens likely represent escapes from cultivation of rabbiteye blueberry, whose genome is formed from that of V.ashei but in many modern cultivars also contains genetic material from one or more other species of V. sect. Cyanococcus. In particular, these plants could be escaped progeny of crosses between V. ashei and V. constablei , another hexaploid species but endemic to the central and southern Appalachians ( Camp 1945; Manos et al. in prep.). Such crosses are known to have been conducted at various blueberry breeding facilities as early as the 1940s ( Brightwell et al. 1949; Ballington et al. 1986, 2006). More recently, cultivars resulting from such crosses, and also those that include V. corymbosum L. or at least germplasm with that name as applied horticulturally, have been released into the market (Galletta & B allington 1996; Ehlenfeldt & Rowland 2015; Ehlenfeldt et al. 2015). Leaf surface features, such as the presence or absence of simple trichomes, in the progeny of such crosses are rarely if ever described in the horticultural literature, rendering it unclear as to whether glabrous leaves specifically result from any such crosses. However, in the wild, both V. constablei and V. corymbosum may have completely glabrous leaves abaxially on occasion, and rabbiteye blueberry cultivars based on V.ashei that also include germplasm from either or both of these species might have been used in the development of these cultivars. The possibility that plants with glabrous leaves abaxially are escaped hybrid cultivars can be tested with genomic analysis.

Wright et al. (2023) hypothesized that plants collected from Prince George’s County, Maryland, were either adventive natural populations of Vaccinium virgatum Aiton (with V. ashei considered a synonym) or escapes from cultivation of rabbiteeye blueberry. If escaped, they likely originated from the nearby Beltsville Agricultural Research Center, where research on “ V.a shei ” was conducted for many years, presumably including crosses with other species. Based on our observations of one of these collections (see below), we consider it likely that these plants and the other presumed escapes cited below include a part of the genome that confers lack of leaf pubescence into the predominant V.a shei genome, thus enabling us to distinguish these plants morphologically from native populations of V.ashei . Some of these specimens have thicker stems and larger leaves than those from the presumed native range of the species, possibly resulting from hybrid vigor in these individuals. Whether or not these escapes are hybrids could be tested by sampling the genomes of presumed escapes and comparing them to those of naturally occurring V. virgatum and other species of V. sect. Cyanococcus. One might then be able to assess whether these plants have genomes from more than one species, and if so, the species involved and their proportional genomic contributions.

Several specimens seen by us that occur outside the native range of Vaccinium ashei as circumscribed here [Alabama, Lee County, P. W.Fritsch2429 ( BRIT, DUKE) from the Auburn University Campus; Georgia, Butts County, C. L. Howel 0076 (NLU, VDB); Georgia, Clarke County, D.Demaree 51237 ( VDB); and Harris County, Georgia, Sweeney 280 ( GA [2 sheets], NLU), 312 ( GA, NLU), and 313 ( GA, NLU), all near Callaway Gardens but apparently not cultivated], have nearly the same level of leaf pubescence as plants in the native range. These plants could either be adventive plants of V.ashei or rabbiteye escapes with characters closer or equivalent to wild-type V.a shei. Other collections made from the same general locality on the Auburn University campus are similar to V.ashei but have glabrous leaves and on this basis we can more confidently recognize them as rabbiteye escapes.

Two specimens resembling Vaccinium ashei collected from Echols County, Georgia [ G. J. Galletta PI 346494 (7-17-15) ( NCU, VSC *) and G. J. Galletta PI 346496 (7-17-19) ( NCU, VSC *)] lie within the natural range of V.ashei but have leaves that are completely glabrous on the abaxial surface, as in the presumed escapes from cultivation outlined above. Their leaves are also notably narrowly obovate, unlike those of natural V.a shei, and possibly indicate derivation from hybridization with V. tenellum or other species of V. sect. Cyanococcus.

One of the samples used for our phylogenetic analysis [North Carolina , Orange County, Occoneechee Mountain State Natural Area, P. S. Manos CY-321 ( BRIT)] has a glabrous abaxial leaf surface and is presumed to lie outside the natural range of Vaccinium ashei . As such, we consider it an escaped rabbiteye cultivar. The phylo-genetic grouping of the CY-321 sample together with the naturally occurring samples of V.ashei can be explained by the likely predominance of the V.ashei genome in rabbiteye cultivars. A follow-up study could attempt to detect in rabbiteye escapes the presence and strength of the genomic signature from V.ashei versus that of other species used in the development of rabbiteye blueberry cultivars.

Additional specimens examined from presumed naturally occurring populations of Vaccinium ashei .— U.S.A. ALABAMA. Baldwin Co.: Alabama Hwy. 12, 30°48′30″N, 87°38′35″W, Dogwood Creek Quad, sandy roadside in full sun, 23 Mar 2006, A. R. Diamond 16191 ( VDB); ca. 10 m W of county road 63, ca. 6 m N of county road 112 (Old Pensacola Road), Bay Minette, 82 m, 30.88946°,−87.73838°,longleaf pine forest, flat area, edge of forest along road, loam with thin leaf litter, dappled shade, shrub 1.8 m t all, leaves whitish green and glaucous beneath, dull whitish bluish green and slightly glaucous above, without stipitate-glandular trichomes on abaxial leaf surface, common, 4 May 2022, P. W.Fritsch2442 ( BRIT, DUKE). Coffee Co.: Kingston, roadside Hwy. 52, 0.8 mi S of Covington County line, shrubs ca. 7 ft tall along edge of woods, 6 May 1997, J. R. MacDonald 10372 (DSC*, DUKE *,KNK*, MISSA *, VDB). Conecuh Co.: Hwy. 29, ca. 5 mi SE of Evergreen at Old Beulah Cemetery, T 5 N, R 11 E, S 36, Red hills area of Alabama, 28 Jun 1985, A. R. Diamond 1095 ( GA); US Hwy. 31 at Travis Bridge,Sepulga River,Red Hills area, T 6 N R 13 E S 30, 17 Aug 1985, A. R. Diamond 1518 ( VDB); ca. 3 m. NE of Hacoda, ca. 5 mi W of Samson, sandy creek bluff,high bush, 29 Mar 1986, R. Kral 73200 ( VDB); W side of Repton by US 84, small seep in pineland, shrubs 1.5–2 m, 17 Mar 2003, R. Kral 93664 ( VDB). Covington Co.: un-numbered dirt road 0.5 mi NE of US Hwy. 331,Covington Wildlife Management Area, T 1 N, R 17 E, S 7, low sandy longleaf pine woods, common shrub, 8 Jul 2001, A. R. Diamond 12504 ( VDB);Brooks School Road (dirt), 0.89 mi N of the Conecuh County line, 294 ft,31.4480000°, –86.678833°, dry sandy soil of a young pine plantation, 12 Jul 2011, A. R. Diamond 22345 ( VDB);along trail ca. 15 m W from SW edge of Buck Pond, ca. 0.25 mi S of jct. Tower Road and Road 34 B, vicinity of Open Pond Recreation Area, Conecuh NF, 69 m, 31.08871°, −86.54340°,longleaf pine forest,along trail near edge of lake,damp salt-and-pepper-colored sand in partial shade, shrub 1.9 m tall, with clumped stems and other scattered clumps nearby,together ca. 10 m diam., stems green beneath, red above,leaves whitish green and glaucous beneath, dull whitish bluish green and glaucous above, immature fruit whitish green and glaucous, ca. 8 × 7 mm, ovoid, common, 3 May 2022, P. W.Fritsch2438 ( BRIT, DUKE);Conecuh NF campgrounds off Ala. 24, pine flatwoods near pond, 20 Mar 1973, G. Gonsoulin s.n. ( VDB); 4.5 mi E Wing on county road 4, high bush, sandy creekbank, 18 Apr 1970, R. Kral 38742 ( GA, VDB) [leaf margins entire].by Ala.54, ca. 4 mi E Florala, high bush of longleaf pine sand ridge, 16 Apr 1970, R. Kral 38626 ( VDB); around Open Pond, Conecuh NF, sandy peat of low areas, shrubs to 2 m, corollas white, 8 Apr 1988, R. Kral 74587 ( VDB); 9.4 mi S Libertyville by Al 55, pine-cypress-tupelo flats, highbush shrub, 26 Jul 1990, R. Kral 78074 ( VDB); around Open Pond, Open Pond Recreation Area,Conecuh NF, SW of Andalusia,high bush of sandy pineland, 28 Mar 1996, R.Kral 85716 ( VDB);Conecuh NF, Hwy. 24, 3.3 mi E of jct. Hwy. 137, roadside, 20 May 1998, J. R. MacDonald 11255 ( VDB); Yellow River, 0.25 mi W of Hwy. 55, woods bordering N bank, occasional shrub to ca. 8 ft tall, 10 Aug 1999, J. R. MacDonald 13342 ( DUKE *, MISS *, MO,USMS*);along county road 10 between Hacoda and Florala,in roadside hedgerow,large shrub, foliage somewhat pruinose-glaucous,reddish tinted, fruit large, black, 21 Jun 1997, R. W. Simmers1997-6 ( VDB); Conecuh NF, ecotone of drain N of county road 4, section 28, E of Bradley, 16 Jun 1998, B. A. Sorrie 9775 ( NCU). Dale Co.: un-numbered dirt road just S of Black’s Creek, 13°34′34″N, 85°38′56″W, Ariton Quad, red clay soil of pine woods, common, 20 Mar 2005, A. R. Diamond 14999 ( VDB). Escambia Co.: E. O.Wilson Nature Adventure Park, ca. 0.14 mi N of parking area, ca. 42 m NW of pond, Brewton, border of flat seepy area, near edge of forest along dirt road, 26 m, 31.10432°, −87.07870°, loam under humus with thick leaf litter, dappled shade, broadleaved forest,shrub 2.4 m tall, 2 clumps ca. 2 m apart, leaves whitish green and glaucous beneath,dull whitish bluish green and glaucous above, very few stipitate-glandular trichomes on abaxial leaf surface, common, 4 May 2022, P. W.Fritsch2441 ( BRIT, DUKE); Conecuh River just W of East Brewton, sandy bottoms, high bush, 29 Mar 1981, R. Kral 66781 ( VDB). Geneva Co.: roadside Debice Road near jct.highway 54, 0.8 mi N of hwy. 10 in Hacoda,occasional shrub with nearby V. arboreum , 28 Apr 1999, J. R. MacDonald 12743 (DSC*, DUKE *, IBE, VDB). Houston Co.: Roadside Hwy. 71, 0.1 mi from Dale County line (Little Choctawhatchee River), lower slope of sandhill woods near low wet woods of river, 24 May 1991, J. R. MacDonald 2002 ( VDB). Pike Co.: W edge parking area of Troy University Arboretum, ca. 250 m S of entrance at Pell Avenue, ca. 120 m E of Park Street, Troy, 156 m, 31.79392°,−85.96235°,mixed pine/broadleaved forest in adjacent seepy area, flat area, open understory, dry beige sand with thin pine leaf litter in dappled shade, shrub 3.0 m t all, 1-stemmed, leaves dull pale green and slightly glaucous beneath,lustrous green and rugose above,immature fruit held upright,whitish green, slightly glaucous, limb dull red, appears to be natural (not cultivated), occasional, 2 May 2022, P. W.Fritsch2433 ( BRIT, DUKE); ca. 31 m N of service road to house at Troy University Arboretum, ca. 160 m S of Pell Avenue, ca. 74 m E of entrance road for the arboretum, Troy, 150 m, 31.79471°, −85.96199°, mixed pine/broadleaved forest,gentle slope toward pond, damp reddish beige clay loam with leaf litter, light shade, shrub 3.7 m tall, ca. 15 clumped stems, candalabraform, others in area shorter than this, leaves pale green beneath, lustrous bluish green and glaucous above,immature fruit whitish green, ovoid, glaucous, occasional, 2 May 2022, P. W.Fritsch2434 ( BRIT, DUKE). FLORIDA. Baker County: near bridge over St. Mary River near Taylor, 19 Feb 1929 and 11 May 1929, W. W.AsheE6759 ( GA). Bay Co.: Flora of St.Andrews Bay, Fla., Plot 203, on sand ridge among hard woods, 22 Mar 1926, L. Haven 3490 ( NY); 1 mi N of Bennett, above Moccasin Creek,edge of hammock, 9 Apr 1966, S. McDaniel 7372 (IBE [2 sheets]);ditto, S. McDaniel 7374 (IBE [2 sheets]);ditto, S. McDaniel 7374 (IBE [2 sheets]); 14 mi W of Wewahitchka, near Sandy Creek, corolla white, 24 Mar 1975, S.McDaniel 19790 (BAYLU*, IBE [2 sheets], MO, VDB). Calhoun Co.: 5 mi NE of Youngstown, edge of bottomland forest, sandy soil, 3 Apr 1969, J. Taylor & C. Taylor 5253 B ( BRIT); Columbia Co.: ca. 5 mi NW of Lake City above Falling Creek, just below falls,moist woods on slope, large shrub 3 m tall,fruits borne erect, R 16 E, T 3 S, S 1, 29 Apr 1966, D. B. Ward 5631 ( GA, IBE, NCU). Duvall Co.: Jacksonville, Fort Caroline National Monument and Timucuan Ecological and Historic Preserve, Fort Caroline, intersection of Monument Road and Fort Caroline Road, 22 Apr 2005, D. E. Giannnasi & W. B. Zomlefer 1234 ( GA). Escambia Co.: Pensacola, 6 Apr 1931, W. W.Ashes.n. ( GA); shrub in old abandoned clay pit opposite 5301 Blue Angel Parkway, oak, privet, and myrtle woods, 2.5 ft tall, 12 Mar 1995, S. Askew 11 (IBE); 1.3 mi NW of Beulah, magnolia-pine creek bottom, 28 Mar 1964, S. McDaniel 4068 (IBE, VDB). Franklin Co.: ca. 0.5 mi N of Bristol by FL Rte.12, 21 Mar 1980, R. K. Godfrey 77516 (IBE); 5 mi E of East Point on Hwy. 98, border of slash pine forest and sandy pine ridge, 5 May 1971, R. L.Lazor 5204 ( GA). Holmes Co.: US 90, 3.5 mi W of junction of route 181, 50 ft pine wooded area around periphery of 3 acre pond on S side of road, plants 1.5–2.1 m tall, leaves serrate, ovate, slightly glaucous beneath, fruit black, 11–12 mm, 26 Jun 1967, G. J. Galletta PI 346511 (7-31-1) ( GA, IBE, NCU, VDB); US 90, 3.5 mi W of junction of route 181, 50 ft pine wooded area around periphery of 3 acre pond on S side of road, plants to 1.5 m tall, 26 Jun 1967, G. J. Galletta PI 346513 (7-31-2) (IBE, NCSC, NCU). Jefferson Co.: just NW of Ward Creek bridge, T 2 N, R 4 E, SE1/4 S 15, along roadcut bank in heavily wooded area, frequent, 19 Mar 1982, L. C. Anderson 5739 ( NCU); 4 mi N of Lamont, 23 Mar 1958, R. Kral 6335 ( GA). Leon Co.: in a grove of sand live oak, Quercus geminata , on sandy ridge just above high water mark of pond, Lost Lake Recreation Area, Apalachicola National Forest, 1 mi W of Springhill Road, 14 Jun 1990, R. K. Godfrey 83888 ( GA); 4.6 mi WSW of center of Tallahassee near Chain Lakes, sandy live oak woods 3 m shrub, fruit immature, slightly glaucous, 30 May 1967, S. McDaniel 9089 (IBE, MISS *, VDB); 3 mi N of Woodville, open ground near mixed woods,corolla white, 28 Mar 1966, R. A. Norris 680 (IBE, RSA *). Liberty Co.: beside Kent Road E of route 65 and NE of Hosford, in moist loamy sand of pine-oak woodland above streamlet, frequent large shrub, leaves sparsely glandular hairy abaxially, 30.41601°N, 84.77009°W, 7 Apr 2016, L. C. Anderson 29520 ( MO, NY); Apalachicola NF, Hwy. 12, across from Forest Road 113 turnoff, 6 ft tall, 30.163848°, −84.971966°, 1 Apr 2019, A. A. Crowl CY-203 ( BRIT);Apalachicola NF, along Hwy. 65, 2–6 ft tall, 30.241688°,−84.868757°, 1 Apr 2019, A. A. Crowl CY-204 ( BRIT); Telogia,along Hwy. 65, 100 m N of Telogia Baptist Church, 7 ft tall, 30.354447°, −84.818547°, 1 Apr 2019, A. A.Crowl CY-206 ( BRIT). Okaloosa Co.: near Valparaiso, “ V. n. sp.,” 1927, W. W.Ashes.n. ( NCU); Lightwood Knob Creek, 8 Jun 1929, W. W.Ashes.n. ( GA, NCU); Tom’s Creek, 8 Jun 1929, W. W.Ashe E 6766 ( GA); Niceville, 7 Apr 1931, W. W.Ashe s.n. ( NCU [6 sheets]); near Niceville, 8 Jun 1929, W. W.AsheE6755 ( GA); near Niceville, 8 Jun 1929, W. W.AsheE6756 ( GA);Niceville,Juniper Bayou, 11 Jun 1929, W. W. Ashe E 6758 ( GA);Camp Walton,Five Mile Bayou, 25 Mar 1927 and 8 Jun 1929, W. W.AsheE6761 ( GA); Eglin Air Force Base, Road 787 and Boiling Creek, just S of Yellow River,Hardwood forest,30.5783°,−86.9000°, 17 m, 8 Aug 2001, A. F.Bradley & J.Stone 912 ( MO); Eglin Air Force Base, Kepner Pond, off Hwy. 85 N of 85-123 junction, pine and oak forest,30.59083°,−86.53750°, 67 m, 7 Aug 2002, A. F. Bradley & J. Stone 1181 ( MO); 1.9 mi W on dirt road off Fla. route 4, E side of small creek spanned by wooden bridge,plant ca. 2.7 m tall, 4 main stems, suckering somewhat, leaves broadly elliptic to ovate, serrate, glandular, glabrous above, yellow-green slightly glaucous on underside, 50–72 mm long, 25–49 mm wide, fruit dark blue with bloom, 11 mm diam., pleasant sweet flavor, not gritty, 26 Jun 1967, G. J.Galletta PI 346502 (7-28-1) (IBE, NCSC, NCU); 1.9 mi W on dirt road off Fla. route 4, E side of small creek spanned by wooden bridge, two plants, to 2.4 m tall, leaves lanceolate to oblanceolate, 26 Jun 1967, G. J. Galletta PI 346504 (7-28-2) (IBE [2 sheets], NCSC, NCU); 2.0 mi W on dirt road off Fla. route 4, oak woods on sandy ridge, plant 1.8 m tall,leaves elliptic,glabrous, and glandular,fruit dark, 9 mm diam., G. J. Galletta PI 346506 (7-29-3), 26 Jun 1976 (IBE, NCSC, NCU); 2.0 mi W on dirt road off Fla. route 4, oak woods on sandy ridge, clonal bush to 2.4 m, stems pubescent,leaves slightly glaucous, pubescent and glandular, fruit dark blue with bloom, 26 Jun 1967, G. J. Galletta PI 346508 (7-29-4) (IBE, NCSC, NCU); 3.2 mi W on dirt road off Fla. route 4, plants 1.8–4.6 m tall, 26 Jun 1967, G. J. Galletta PI 346510 (7-30) (IBE, NCSC, NCU);Yellow River hills, W of Laurel Hill, open oak-pine woods, 2 Aug 1961, R. K. Godfrey 61280 ( BRIT); open upland pine-deciduous woodland, 2 mi W of Laurel Hill, 26 Jun 1964, R. K. Godfrey 64380 (IBE); 2 mi N of Niceville along route 85, titi bottom, buds pink, 2 Apr 1966, R. Kral 26142 ( BRIT, GA, IBE, VDB); 2 mi E of Holt, abundant along roadside, 2.5 m tall, 27 Mar 1964, S. McDaniel 4030 (BAYLU*,IBE [2 sheets], VDB [2 sheets]); 2 mi E of Crestview near Adams Creek, at base of hillside, 1.5 m tall, fruit black, 31 Jul 1964, S. McDaniel 4957 (IBE); Eglin Air Force Base, along the Shoal River at Gin Hole Landing, ca. 6.5 mi W of Florida State Rte. 85 off of Air Force Base Road 211, 40 m, 30°40′59″N, 86°38′58″W, shrub 2 m tall,fruits green, 2 Jun 1998, J. S. Miller & J. Stone 9458 ( BRIT, MO); Eglin Air Force Base, 7.5 mi W of Florida State route 85 on Air Force Base route 211, 40 m, 30°39′51″N, 86°41′07″W, shrub 2 m tall, fruits green, 5 Jun 1998, J. S. Miller et al.9543 ( MO, BRIT);Crestview, Mar 1919, M. A. Sapp E 7036 ( GA);sandy hills near Crestview, 7 Apr 1925, J. K.Small & E. T.Wherry s.n. ( NY [2 sheets]); ditto, 11716 ( NY [2 sheets]). Santa Rosa Co.: near Milton, 1 Jul 1931, W. W.Ashe s.n. ( NCU); river swamp, 1 mi W of Milton, 5 Aug 1954, E. S. Ford & L. E. Arnold 4154 ( GA); just W of Wallace in Escambia River bottoms, high bush, 11 Apr 1964, R. Kral 19688 (SMU, VDB); 1 mi E of Harold, longleaf pine slope above Chamaecyparis thicket, ca. 4 m tall, 24 May 1964, S. McDaniel 4596 (IBE); ditto, much branched shrub ca. 3 m tall, 24 May 1964, S. McDaniel 4600 (IBE); 1.3 mi N of East Bay River on Fla.87, 22 May 1976, J. C.Solomon 2310 ( MO); Blackwater River SF, Forest Roads 1, 21, and 28, plants in mesic to moist pine slopes with Ilex coriacea, Osmanthus americanus, Cyrilla racemiflora , Vaccinium arboreum ,multi-stem shrubs to 7 ft tall, frequent, 12 and 13 Jun 1998, B. A. Sorrie 9765 ( NCU). Taylor Co.: Salem, slash pine-palmetto scrub, high bush, 17 Mar 1976, R. Kral 57335 ( VDB). Wakulla Co.: end of Purify Bay Road, edge of St.Mark’s National Wildlife Refuge, 8 ft tall, 30.090715°, −84.371436°, 2 Apr 2019, A. A. Crowl CY-210 ( BRIT);by forest road leading to East River from near the entrance to St.Marks Wildlife Refuge, Lighthouse Road,pine flatwoods,plant forming a dense thicket ca. 5 m across, stems to 2 m tall, leaves slightly glaucous beneath, 24 May 1978, R. K. Godfrey 76513 ( VDB). Walton Co.: Rocky Run, 23 Mar 1927 and 7 Jun 1929, W. W. AsheE6763 ( GA); Alaqua Creek, 26 Mar 1927 and 7 Jun 1929, W. W.AsheE6764 ( GA);along Rte. 183 near Knox Hill Baptist Church, on edge of woods, shrub to ca. 1.5 m t all, 19 Apr 1982, D. S. Correll & H. B. Correll 53755 ( NY); Natural Bridge Road (dirt), 0.09 mi N of Walton County Hwy. 181,192 ft,30.990500°,−86.207927°,sandy wet drain, 3Apr2014, A. R. Diamond 24926 ( VDB);Villa Tasso, 1 mi W of Choctaw Beach in very sandy pine-live oak-myrtle oak woods, 3 M ay 1967, S. McDaniel 8858 ( MISS *). Washington Co.: 1 m i W of Hinsons Crossroads, dry open woodland,shrub ca. 2 m high, 3 Apr 1958, R. K. Godfrey 56372 ( GA,SMU); 1 mi W of Millers Ferry, dry open woodland, shrub ca. 1 m high, 3 Apr 1958, R. K. Godfrey 56373 (SMU); W side of Orange Hill, 7 mi SE of Chipley, T 3 N R 13 W S 2, 16 Oct 1972, D. B. Ward 8480 (IBE). GEORGIA. Camden Co.: Little Satilla River directly E of Waverly, 0.8 mi SW of Camden-Glynn county line, 26 Jun 1952, W. H. Duncan 13792 ( GA). Charlton Co.: Okefenokee National Wildlife Refuge, Stephen C Foster State Park, along boardwalk, 30.82798°, −82.36294°, 5 ft tall, 27 Mar 2021, A. A. Crowl CY-376 ( BRIT);Okefenokee Swamp at end of route 177,plant 1.8 m tall, 24 May 1967, G. J. Galletta PI 346498 (7-20-1) ( VSC *, NCSC, NCU); Okefenokee Swamp at end of route 177, plant 1.8 m t all, 24 May 1967, G. J. Galletta PI 346499 (7-20-6) (GAS [2 sheets]*, NCU); Okefinokee Swamp,Chesser’s Is., hammock, 6–7 ft high, bark smooth, light gray, 25 May 1930, J. S. Harper 135a ( NY); near Toledo, 28 Jun 1930, H. M. McKay s.n. ( GA,IBE). Echols Co.: 1 mi S of Lanier County line on US 29, flat open woodland, G. J. Galletta PI 346491 (7-17-6) ( MO, NCSC, NCU, VSC *); 1 mi S of Lanier County line on US 29, flat open woodland, plant 2.1 m tall, leaves pubescent, serrate, obovate, to 41 mm long by 24 mm wide, G. J. Galletta PI 346493 (7-17-14) ( NCSC, NCU, VSC *); 1 mi S of Lanier County line on US 29, flat open woodland, plant to 2.4 m tall, leaves pubescent, serrate, obovate, 46 mm long and 22 mm wide, G. J. Galletta PI 346497 (7-17-13)( VSC *). Grady Co.: high bush, sandy silt of bottoms, Tired Creek,by GA 93, S of bridge, S of Cairo, 15 Mar 2001, R. Kral 90647 ( VDB);just E of GA 111, N side of Grady County Sheriff’s Dept. bldgs., sandy loam of pine pocosin remnant, shrubs 1.5–2 m, 1 May 2001, R. Kral 90941 B ( VDB) [leaf margins entire and some leaves with secondary veins pubescent abaxially]. LOUISIANA. Washington Parish: 2 mi N of Sun, at edge of woods,shrub 7 ft tall, 12 Jun 1966, J. W. Thieret 23484 ( BRIT). MISSISSIPPI. Forrest Co.: ca. 1 mi W of Hwy. 49 N of Hattiesburg (ca. 2 mi NW of 49-59 intersection), dry mixed woods, plants ca. 6–7 ft tall, 8 Oct 1972, K. E. Rogers 8819 (IBE, SMU). Hancock Co.: Bottoms, Caesar, 8 ft tall, 18 Jun 1953, D. Demaree 33471 ( GH, USF *); 2 mi N of Kiln, cut-over longleaf pine woods, leaves and hypanthium glaucous, 8 Apr 1962, S. McDaniel 2914 (IBE). Harrison Co.: Lyman, up to 3 m tall, corollas pink, 19 Mar 1955, G. R. Cooley & A. S. Pease 3198 ( MISSA *). Jackson Co.: S side of N part of Green Loop of Fontainebleau Trail, ca. 50 m W of N end of Red Loop Trail, ca. 0.28 mi NE of trail head, ca. 0.22 mi E of Hanshaw Road, Mississippi Sandhill Crane National Wildlife Refuge, 6 m, 30.40042°,−88.75364°,longleaf pine forest, along trail in flat longleaf pine savannah, woody undergrowth regenerating in most places, grayish white sand in dappled shade, shrub 1.2 m tall, clumped clone ca. 4 m diam.(extent),2nd-year branchlets olive yellowish green, leaves whitish green beneath, green and slightly glaucous above, 2 clumps of plants seen, 24 Apr 2022, P. W.Fritsch2394 ( BRIT, DUKE); N side of N part of Green Loop of Fontainebleau Trail, ca. 115 m W of N end of Red Loop Trail, ca. 0.27 mi NE of trail head, ca. 0.19 mi E of Hanshaw Road, Mississippi Sandhill Crane National Wildlife Refuge, 6 m, 30.40061°,−88.75436°, longleaf pine forest,along trail in flat longleaf pine savannah,woody undergrowth regenerating in most places,grayish white sand in dappled shade, shrub 1.9 m tall, several stems clumped and then 4 more scattered within ca. 2 m diam. area, immature fruit not notably upright, whitish green, glaucous,with red limb,2 clumps of plants seen, 24 Apr 2022, P. W.Fritsch2395 ( BRIT, DUKE). Pearl River Co.: Picayune, Hobolochitto Creek near US route 11, low magnolia-spruce pine woods, plant 1.5 m tall, 28 Mar 1982, S. McDaniel 25871 (IBE [4 sheets], MISS *, VDB). Stone Co. ca. 4 mi SE of Wiggins, S 34/27,roadside secondary area, ca. 1 m, sterile,apparently spread from cultivation, 30 Jul 1981, S. McDaniel 25255 (IBE; to us, the evidence that this is a cultivated escape is unclear;it has the pubescent midvein abaxially of V.ashei and falls within its apparent natural range); ca. 2 mi N of jct. state routes 15 and 26, well-drained mixed woods, 30 Jul 1981, S. McDaniel 25256 (IBE).

Specimens cited by Reade (1931) as Vaccinium ashei in the protologue but not found online or at herbaria searched.— U.S. A. FLORIDA. Escambia Co.: near Pensacola, 27 Mar 1927, W. W.Ashes.n. Okaloosa Co.: Niceville, Juniper Bayou, 10 Jun 1929, W. W.Ashes.n. Santa Rosa Co.: Hollyhead, 8 Jun 1929, W. W.Ashes.n. MISSISSIPPI. Pearl River Co.: McNeill, 8 May 1930, W. W.Ashes.n.; near Poplarville, 8 May 1930, W. W.Ashes.n.

Specimens of V. ashei of uncertain provenance.— U.S. A. FLORIDA? Duval Co.?: near Jacksonville,dry oak woods, 20 Feb 1894, A. H. Curtiss 4522 C ( NY); near Jacksonville, low thickets, 3–8 ft high, 21 Feb and 10 May 1894 (“from different localities”), A. H. Curtiss 4522 ( MO, NY).These collections may have been made in any one of various counties near Jacksonville, or even in Charlton Co., Georgia, and thus are not mapped here.

Specimens examined of presumed adventive plants of Vaccinium ashei and rabbiteye blueberries escaped from cultivation.— U.S.A. ALABAMA. Geneva Co.: high bush of edge of cypress-gum pind by Co.4, 6.9 mi W jct. AL 167, 17 Jul 1992, R. Kral 81170 ( VDB). Lee Co.: estate of Dr. H. P.Orr-Opelika, 15 Jul 1970, T. A. Burrough 058 ( VDB);in the Forestry Plot off of Donahue Ave.near the lake in Auburn, 1 Apr 1973, B. Campbell 030 ( VDB); along trail starting ca. 100 m E o f boardwalk trail at Hog Pond, ca, 64 m S o f Woodfield Drive, Auburn University, 194 m, 32.58796°, −85.49309°,mixed pine/broadleaved forest, along trail at edge of seepy area on very gentle slope,light shade, dry loam with thin leaf litter, occasional, shrub 3.0 m t all, multi-stemmed in 2 clumps, with other stems near these, one branchlet a 2.1-m-long erect “sucker” off the main stem, 2nd-year branchlets olive green, current-year branchlets green, leaves whitish green and slightly glaucous beneath, green and slightly glaucous above,immature fruit held upright,light green and slightly glaucous with red limb, 2 May 2022, P. W.Fritsch2429 ( BRIT, DUKE); Auburn University Forestry Plot covering an area from the hog pond circling around to Donahue Drive, small tree growing in groups, scattered throughout woods, 29 Jun 1971, C. J. Hayden III 067 ( VDB). GEORGIA. Butts Co.: SE of Jackson, off GA Hwy. 42, Indian Springs State Park,along Hopoetnyeloholo Creek below miniature golf course, shrub 1.5 m tall, 12 Apr 1987, C. L. Howel 0076 ( GA, NLU, VDB). Charlton Co.: Scrub, sandy soil, Floyd’s Island, 29 Sep 1959, E. Cypert 236 ( GA). Clarke Co.: P. O. Athens,low ridge, 5 ft tall, 750 ft, 6 Sep 1964, D.Demaree 51237 ( VDB). Echols Co.: 1 mi S of Lanier-Echols County line beside trash barrel on US 129, and also at Norsworthy farm ca. 1 mi further S on same road, flat open woodland, obovate leaves, 24 May 1967, G. J. Galletta PI 346490 (7-17-2) ( GA);ditto, G. J. Galletta PI 346492 (7-17-9) ( MO, NCU,PAC*, VSC *);ditto, plant 1.8–2.1 m tall, leaves glaucous, serrate, oblanceolate, to 50 mm long by 18 mm wide,fruit very light blue, G. J. Galletta PI 346494 (7-17-15) ( NCU, VSC *);ditto, G. J. Galletta PI 346496 (7-17-19) ( NCU, VSC *). Elbert Co.: NE of Elberton,dirt road 115off Hwy. 17, edge of deciduous woods, 9 Apr 1977, N. Craft Coile 446 ( GA). Harris Co.: Beaver Pond, 16 Jul 1998, Crook 489 ( GA); dam near entrance to Laurel Springs Dr., Callaway Gardens at Pine Mountain, 13 Jul 1972, S. B. Jones 22155 ( GA);flora of Calloway Gardens, ca, 100 yds. S of the entrance to Dog 7 on Dog 7, common, successional forest, 6 Apr 1999, Sweeney 280 ( GA [2 sheets], NLU); flora of Calloway Gardens, Callaway Airport, S side of runway, common margin of disturbed field, 6 Apr 1999, Sweeney 311 ( GA);ditto, Sweeney 312 ( GA,NLU);ditto, Sweeney 313 ( GA,NLU). Liberty Co.: low sandy ridge, 11 mi E of Taylor, 27 Apr 1940, W. H. Duncan 2238 ( GA). Tattnall Co.: just SW of Reidsville city limits, 26 May 2010, L. A. Jackson 29 ( GA). MARYLAND. Prince Georges Co.: Drainage of Patuxent River, just below the NE end of SW-to-NE runway of the abandoned Beltsville Airport, 4 km NW of Bowie, 28 Sep 1999, M. T. Strong 2231 (GMUF GMUF-0023458).Other specimens cited in Wright et al. 2023 from this county have not been seen by us but we presume that they are also adventive or escapes from cultivation. MISSISSIPPI. Jasper Co.: T 4 N R 10 E S 3 and 4 behind wildlife management area headquarters bldg., mature pine forest fairly open canopy mixed oak and hickory understory, periodic prescribed fire use, common open and disturbed areas, 16 Jun 1988, D. T. Carraway 80 (IBE); ditto, D. T. Carraway 81 ( VDB; perhaps a mislabeling of D. T. Carraway 80 because the label data has an identification of Cyperus pseudovegetus Steud. NORTH CAROLINA. Durham Co.: Duke Forest off of Gate 10 entrance, 36.022586°,−78.982696°, 31 Mar 2016, P. S. Manos CY-010 ( BRIT). Harnett Co.: pocosin,Spout Springs, 22 Apr 1970, R.Wyatt 103 ( GA). Orange Co.: Occoneechee Mountain State Natural Area, intersection of Brown Elfin Knob Trail and Mountain Loop Trail,36.063783°, −79.113196°, 7 Sep 2019, P. S. Manos CY-321 ( BRIT). Moore Co.: beside trail on W side Heron’s Lake, Whispering Pines, edge of woods with Quercus alba , Q. nigra , Acer rubrum , Pinus taeda , Ilex opaca , leaves bluegreen above, glaucous beneath, 35.270000, −79.372222, 17 May 2023, B. A.Sorrie 14145 ( NCU). SOUTH CAROLINA. Barnwell Co.: Savannah River Site:along Patterson Mill Road, 10 Jun 1989, N. Coile 5348 ( GA). McCormick Co.: Upland pine woods ca. 1 mi E of Modoc, 17 Apr 1970, W. H. Duncan 23376 ( GA). TEXAS. Smith Co.: ca. 100 m SW of Blackhawk Creek and ca. 400 m (direct) from end of County Road 2235 E of TX-110, property of Ron & Ruth Loper, 115 m, 32.18574°, −95.18666°, mixed pine/broadleaf forest, in humus and clay over sand in sun, shrub two-stemmed at base, 2.4 m tall, current-year branchlets, leaves, and fruit pale glaucous, apparent escape from cultivation with possible parent Fritsch 2260,only one plant, 21 Apr 2021, P. W.Fritsch2259 ( BRIT, DUKE).