Semple, J.C. 2012. Typification of Sofidago graciffima (Asteraceae: Astereae) and application of the name. Phytoneuron 2012-107: 1-10. Published 5 December 2012. ISSN 2153 733X TYPIFICATION OF SOLIDAGO GRACILLMA (ASTERACEAE: ASTEREAE) AND APPLICATION OF THE NAME JOHN C. SEMPLE Department of Biology University of Waterloo Waterloo, Ontario Canada N2L 3G1 jesemple@uwaterloo.ca ABSTRACT Variations in application of the name Solidago gracillima have compounded confusion over what to include in synonymy under the name. It is regarded here as a species distinct from all those previously included in its synonymy. The holotype shows a diagnostically large inflorescence a few long branches; probable isotypes are variants with smaller, compact inflorescences without elongated lower branches. Morphological features of S. gracillima are discussed and compared with other species in Solidago subsect. Maritimae. KEY WORDS: Solidago gracillima, Solidago austrina, Solidago stricta, Solidago subsect. Maritimae Solidago gracillima Torr. & A. Gray is a goldenrod species native to the outer edge of the piedmount and the Fall Line counties of South Carolina, Georgia, Florida, and possibly North Carolina. It is a member of Solidago subsect. Maritimae (Torr. & A. Gray) G.L. Nesom, a group of bog., marsh, and seasonally wetland goldenrods with lower stem and basal rosette leaves that have petioles sheathing the stem (Semple & Cook 2006). It has been treated as a relatively narrowly defined species distinct from S. stricta Ait. and S. austrina Small (Small 1903; Cronquist 1968), as a weakly distinct species "perhaps not specifically distinct" from S. stricta" (Radford et al. 1968), as a broadly defined species including S. austrina and S. simulans Fern. (Jones & Coile 1988) and S. perlonga Fern. (Cronquist 1980), or as a subspecies within S. stricta Ait. (Semple & Cook 2006, including S. austrina Small). The confusion is the result of 1) a lack of understanding about inflorescence variation in subsect. Maritimae, 2) how many species should be recognized within subsect. Maritimae, and 3) how best to distinguish the species that are recognized with the subsection. Solidago gracillima Torr. & A. Gray, Fl. N. Amer. 2(2): 215. 1842. Solidago stricta Ait. subsp. gracillima (Torr. & Gray) Semple, Sida 20: 1615. 2003. TYPE: USA. Florida. "Middle," Dr. Chapman s.n. (holotype: NY!, Fig. 1; probable isotypes: KEW 2 sheets!, NY 3 sheets!). The holotype, as recognized here, may be the only collection at NY seen by Torrey and Gray and is the only specimen marked “S. gracillima n. sp.” on a Torr. & Gray, Flora N. Amer. label. It also is the only one with an original, printed annotation of "Syn. Fl. N. Amer." Several other probable duplicates of this collection are now at NY but were originally at Columbia College Herbarium (Fig. 2), Hamilton College Herbarium (Fig. 3), and Columbia University Herbarium (Fig. 4). The latter (as well as the holotype) is annotated as “Solidago gracillima TYPE K.M.” (K. MacKenzie). The exact collection locality in Florida is not indicated on any of the Chapman s.n. specimens. The species occurs in Panhandle Florida. The Atlas of Florida Vascular Plants (Wunderlin & Hansen 2012) reports collections from Bay, Franklin, Leon, and Wakulla counties, but only collections from Leon County have been seen from Florida by the author. p. F Solidi illima Torr. & Gray,, Fl. N. Amer. 2(2): 215. 1842. Solidago stricta Ait. subsp. gracillima (Torr. & Gray) Semple. SYNTYPES: U.S.A ‘ FLORIDA. "middle," Dr. Chapman s.n. (LT [to be designated KEW! 1 NY(3)! 5 John C. Semple, WAT 22 Oct 2012 University of Waterloo TYPE OF cracillima Torrey & Gr¢ mer, 2: 215. 182. ESolidago Fl. N dup. ver. M. A etter 7/1993 | Roger Boettcher 197% 4 * Louisiana State University, Baton Rouge Y Grae hepa x») Ae P uthern Unite States, Lop la he form SYN. FL. N. AMER. iii i 2 ‘Tour. & Guay, Fiona, N, Ann. hy —, | Figure 1. Holotype of Solidago gracillima Torr. & A. Gray, Chapman s.n. (NY). a ‘Solidago gracttia or. 4 emp Det John ©. Sample 2012 Wat LUnwersy of Waterco Joba. Serle, WAT 2£0st20)2 —niversty oF Waterco Solidago geaciitima Tare. 8 A, Gray N / 2-KEW Yor POTANICAL ® GaapeNn Be. pe 2 ( Figure 2. Probable isotype of Solidago gracillima, Chapman s.n. (NY ex Columbia College Herb.). ISOLECTOTYPE: Solidago gracillima Torr. & A.G N. Amer. 2(2): 215. 1842. Solidago stricta Ait. s tr. & Gray) Semple. SYNTYPES Je," Dr. Chapman s.n. (LT [to be designated Semy neuron}: NY! ILTs:, KEW! (several sheets), NY(3)! John C. Semple, WAT 22 Oct 2012 niversity of W prohahle TYPE OF Solidago gracillima Torrey & Gray Flora N. Amer. 2: 215. 1842. ver. M. A. Wetter 07/198 Sat olag « gractlimes LPG nll! Aus, Chae / 4 / ON frAn err Figure 3. Probable isotype of Solidago gracillima, Chapman s.n. (NY ex Hamilton College Herb.). ya greet n= W YOR é fee = ~J < \ ITANIC4L DEN _— | ISOLECTOTYPE: ope be ge rion orr. & A f N. Amer. 2(2); 215. 1842. Solidago ns er g is ( Torr, & y) Semple. SYNT as I \. # "middle," Dr res n. (LT [t lesignated n ¢ Phytoneuron]: NY! ILTs:, KEW! (several sheets), NY( § F John C. Semple, WAT 220 University of \ i a Solidago gract1lima T.6 ( Figure 4. Probable isotype of Solidago gracillima, Chapman s.n. (NY ex Herb. Columbia University). Type material of Solidago gracillima (Chapman s.n. NY, Florida) includes large inflores- cences that are very open with a few long branches (holotype NY, Fig. 1; isotype NY, Fig. 2) and smaller compact inflorescences without elongated lower branches (isotypes NY, Figs. 3-4). The same individual can produce different shoots with either large or small inflorescences. Terminal and elongated lateral branches have secund heads on distally arching stems. In comparison, inflorescences of S. stricta are elongate and narrow with short usually ascending branches (Fig. SB). The apex is erect and not secund, unless the entire stem is arching as the infloresence develops. Then, the entire inflorescence may be one-sided with branches growing upward in the same direction. If the secund inflorescence of S. gracillima is pressed and dried so that the arching is flatted out, then a small inflorescence could be easily confused as that of S. stricta. Inflorescences of S. austrina when large have ascending spreading elongated lower branches (Fig. 5A); the apex is often slightly secund. Small inflorescences can be like those of S. stricta. Inflorescences of S. perlonga are similar to S. austrina, but the lower elongated branches are longer and more widely spaced on the stem (Fig. SC). Smaller inflorescences even on the same clone can be similar to those S. stricta (Fig. 5C). Other species in the subsection have slightly different to obviously different inflorescences. Those of Solidago uliginosa (Fig. 5D) have short ascending branches and are club-shaped but can be similar to those of S. simulans (Fig. SE) or even S. austrina. Inflorescences of S. mexicana (Fig. 5F) and S. sempervirens (Fig.. 5G) are more secund-pyramidal in general shape. Those of S. mexicana have small stem leaves near and into the inflorescence, while those of S. sempervirens usually have large leaves just below and into the inflorescence. Those of S. mexicana can be similar to S. stricta, if the secund aspect to the apex is not strongly developed. In all species, size of the inflorescence is critical in determining branching pattern. Strong apical dominance requires lower branches to be relatively distant before the lower branches can elongate into diagnostic patterns. The tendency for botanists to collect specimens that fit on herbarium sheets, i.¢., mid to small individual shoots, has resulted in numerous specimens with small inflorescences with non-diagnostic features. This has led to many misidentifications and repeated errors in the literature about the distribution of individual species. Work is underway to accurately map the distributions of all species in the subsection. Rosette leaves and lower stem leaves of Solidago gracillima are shallowly serrate distally or along much of the margin (Fig. 6). The holotype includes several lower stem leaf fragments that are clearly serrate (Fig. 6A). One of the probable isotypes includes a rosette with rounded-spatulate to acute-lanceolate leaves that are long petiolate and serrate distally (Fig 5D). In the field, lower stem leaves can be similar to the acute, lanceolate rosette leaves and these maintain those traits in cultivated transplants (Fig. 6 B-C respectively, Semple & Semple 11834). A few herbarium specimens have larger broader rosette leaves, but under cultivation in a growth chamber such leaves are very robust for the species (Fig. 6 E; seedling rosette leaves, Anderson 25350 FSU). Basal leaves of S. austrina are similarly serrate but usually larger and oblanceolate, based on observations of lower stem and rosette leaves of herbarium specimens from BRIT, FSU, GA, GH, MO, MT, NY, NCU, USCH, and USF (Thiers, continually undated) and field observations of wild plants in North Carolina, South Carolina, Georgia, and Tennessee. Rosette leaves of S. stricta are variable in shape from linear oblanceolate to lanceolate to broadly obovate. However, these are never serrate although sometimes crenate. Figure 5. Inflorescence variation in Solidago subsect. Maritimae. A. S austrina, Semple & Semple 11203, Alabama. B. S. stricta, Semple 11777, South Carolina. C. S. perlonga, Semple 11824, New Jersey. D. S. uliginosa, Semple 11837, Michgan. E. S. simulans, Semple 11588, North Carolina. F. S. mexicana, Semple 11651, North Carolina. G. S. sempervirens, Nova Scotia. sate ss ee s Figure 6. Stem and rosette leaves of Solidago gracillima. A. Lower stem leaf fragment, holotype (see Fig. 1); scale bar = 1 cm. B-C. Lower stem leaves of Semple & Semple 11834. B. Wild plant growing out of vertical road embankment. C. Cultivated plant grown from rootstock transplanted to WAT. D. Basal rosette leaves of isolectotype (see Fig. 2). E. Growth chamber grown base rosette of seedling from Anderson 25350 (FSU). Wild populations of Solidago gracillima have been seen in South Carolina and southwestern Georgia. A small population encountered in Barnwell Co., South Carolina, in 1981 Gemple & Suripto 9814) grew at the top of a bank around a shallow pool in a mixed pine and broadleaf woods. Several populations sampled in southwestern Georgia in 2010 were in habitats that could potentially be wetter in the early season, but were dry in early September. Semple & Semple 11834 (WAT) grew ona sandy clay ridge and on the face of a very steep road cut in an area of mature pine forest and pine plantations (Fig. 7A). Semple & Semple 11836 (WAT) grew in drier, sandy soil at the top of a roadside ditch, while a few individuals of S. austrina grew at the bottom of the ditch in wet sandy and mucky soil (Fig. 7B). All collections of S. austrina seen in North Carolina, South Carolina, Georgia, and Tennessee grew in ditches, seeps, wet prairie, and heads of tributaries in usually wetter soils. Solidago g tai i Figure 7. Habitats of Solidago gracillima. A. Semple & Semple 11834, SE of Quitman, Brooks Co., Georgia. B. Semple & Semple 11836, SE of Cairo, Grady Co., Georgia. 10 Overall, the habitats of Solidago gracillima are likely seasonally drier than those of S. austrina. Both species can be locally common, but S. gracillima is generally less common. I conclude that the tendency to grow in habitats that change from wet to dry over the season is the reason many herbarium collections lack mid and lower stem leaves. The larger leaves become ecologically unsuited to the drier late season conditions and are dropped by the plant. In contrast, most species of subsect. Maritimae grow in habitats that are wet to very moist throughout the season. These tend to have lower stem and rosette leaves present at the time of flowering. ACKNOWLEDGEMENTS This work was supported by a Natural Sciences and Engineering Research Council of Canada Discovery Grants to the author. Joan Venn is thanked for her curatorial assistance with loans. The following herbaria are thanked for loaning specimens of Solidago subsect. Maritimae: BRIT, FSU, GA, GH, MEXU, MO, MT, NY, NCU, TEX, USCH, and USF. The following students assisted in recording location data and collecting morphological data on specimens of subsect. Maritimae: Mahmoud ElI-Sweisi, Andrew Lam, Yunfei Ma, Hammad Rahman, Andrea Ranson, Troina Shea, Marion Sorour, and Lan Tong. LITERATURE CITED Cronquist, A. 1968. Solidago L. Pp. 413-438, in H.A. Gleason (ed.). The New Britton and Brown Illustrated Flora of the Northeastern United States and Adjacent Canada. Hafner Pub. Co., New York. Cronquist, A. 1980. Vascular Flora of the Southeastern United States. I. Asteraceae. Univ. North Carolina Press, Chapel Hill. Jones, S. B. and N.C. Coile. 1988. The Distribution of the Vascular Flora of Georgia. Department of Botany, Univ. of Georgia. Athens. Radford, A-E., H.E. Ahles, and C.R. Bell. 1968. Manual of the Vascular Flora of the Carolinas. Univ. of North Carolina Press, Chapel Hill. Semple, J.C. and RE. Cook. 2006. Solidago Linnaeus. Pp. 107-166, in Flora North America Editorial Committee (eds.). Flora of North America. Vol. 20. Asteraceae, Part 2. Astereae and Senecioneae. Oxford Univ. Press, New York. Small, .K. 1903. Flora of the Southeastern United States. Published by the author, New York. Thiers, B. [continuously updated]. Index Herbariorum: A global directory of public herbaria and associated staff. Virtual Herbarium, New York Botanical Garden. http://sciweb.nybg.org/science2/IndexHerbariorum .asp Wunderlin, RP. and B.F. Hansen. 2012. Atlas of Florida Vascular Plants. [S.M. Landry and K.N. Campbell (application development), Florida Center for Community Design and Research] Inst. for Systematic Botany, Univ. of South Florida, Tampa.